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Scottish Natural Heritage Research Report No. 882

South Arran MPA diver survey of maerl beds, kelp and seaweed communities on sublittoral sediment, and seagrass beds 2014

R E S E A R C H R E P O R T

Research Report No. 882

South Arran MPA diver survey of maerl

beds, kelp and seaweed communities on

sublittoral sediment, and seagrass beds

2014

For further information on this report please contact:

Lisa Kamphausen Scottish Natural Heritage Great Glen House Leachkin Road INVERNESS IV3 8NW Telephone: 01463 725014 E-mail: [email protected]

This report should be quoted as:

Mercer, T., Kamphausen, L., Moore, J., Bunker, F., Archer Thompson, J. & Howson, C. 2018. South Arran MPA diver survey of maerl beds, kelp and seaweed communities on sublittoral sediment, and seagrass beds 2014. Scottish Natural Heritage Research Report No. 882.

This report, or any part of it, should not be reproduced without the permission of Scottish Natural Heritage. This permission will not be withheld unreasonably. The views expressed by the author(s) of this report should not be taken as the views and policies of Scottish Natural Heritage.

© Scottish Natural Heritage 2018.

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South Arran MPA diver survey of maerl beds, kelp and seaweed communities on sublittoral sediment, and

seagrass beds 2014

Research Report No. 882 Project No: 015165 Contractor: Aquatic Survey & Monitoring Ltd. Year of publication: 2018 Keywords

MPA; South Arran; maerl beds; kelp and seaweed communities on sublittoral sediment, seagrass beds; survey; diving Background

The South Arran Marine Protected Area was designated by the Scottish Government as one of 33 new Nature Conservation Marine Protected Areas (MPAs) in 2014. A series of surveys were conducted around that time to collate the evidence base for the MPA and to establish a baseline against which to assess the effectiveness of any new management measures. The results of one of these surveys are presented here - a diving survey which collected detailed biological information on three of the MPA’s protected features: ‘maerl beds’, ‘kelp and seaweed communities on sublittoral sediment’, and ‘seagrass beds’. The results formed part of the documentation underpinning the MPA designation process. The ‘maerl beds’ and ‘kelp and seaweed communities on sublittoral sediment’ features encompassed within the South Arran MPA have been exposed to demersal fishing activity for many years. Maerl (the collective term for several species of calcified red seaweed which in their free-living form and under favourable conditions can create extensive beds) is particularly sensitive to pressures arising from bottom-contacting fishing activity. Maerl habitats studied to date within the MPA have generally been considered to be in poor condition with a very low proportion of live maerl cover. The maerl beds feature within the South Arran MPA has therefore been assigned a ‘recover’ conservation objective. A threshold of 5% cover of live maerl has been adopted for maerl biotope recognition within the MPA to align with previous studies (Moore, 2014). On this basis, beds of living maerl are currently believed to be confined to Lamlash Bay and to small pockets off the south coast (around Pladda and the Iron Rock Ledges), in areas of formerly low levels of fishing pressure. More extensive areas that incorporate clusters of records of sparse living maerl and relatively unbroken dead maerl have also been identified (including off the south of Holy Isle). These ‘maerl recovery areas’, which are believed to represent areas of historically richer maerl, appear to offer the most suitable targets for management action for this feature and were included within broader management proposals consulted upon at the end of 2014 (Marine Scotland, 2014).

RESEARCH REPORT

Summary

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The dive survey presented here was designed to sample different maerl beds (areas known to support living maerl exceeding 5% cover) and maerl gravel habitats (where the proportion of live maerl material was less than 5% cover) in comparable shallow infralittoral settings across the MPA. The three maerl gravel sites studied were part of the ‘kelp and seaweed communities on sublittoral sediment’ protected feature. The aims of the work were to determine the respective levels of biological diversity in these habitats and to investigate the differences in associated communities with differing proportions of live maerl. Sampling locations were established inside and outside the Lamlash Bay No Take Zone and within maerl recovery areas off Pladda and to the south of Holy Isle. Together with the other work conducted to establish a baseline of condition in the South Arran MPA, these data will aid the assessment of the effectiveness of the recently introduced management measures, and chart recovery of relevant degraded maerl habitats. Main findings

Two maerl beds in the South Arran MPA were surveyed in detail by divers for the first time. With 29% and 76% live maerl coverage; these beds contained by far the highest proportions of live maerl of any extant maerl beds in the Clyde. These records show that Clyde maerl beds can comprise a continuous layer of well-developed live fragments. Both beds had a relatively rich infaunal community and rich epibiota.

One of the two newly recorded maerl beds, located in Lamlash Bay, was of the biotope SS.SMp.Mrl.Lgla - Lithothamnion glaciale maerl beds in tide-swept variable salinity infralittoral gravel. The bed was dominated by fine twigs of L. glaciale. This is the first instance of a record of this biotope in the Clyde. All other maerl bed records so far were of biotopes based on a different maerl species (Phymatolithon calcareum - SS.SMp.Mrl.Pcal). A detailed record of species associated with this new biotope was created.

The other rich maerl bed recorded during this survey, west of the Isle of Pladda, was of the biotope SS.SMp.Mrl.Pcal.R - Phymatolithon calcareum maerl beds with red seaweeds in shallow infralittoral clean gravel or coarse sand. The Phymatolithon calcareum on this much more exposed bed formed large medallions.

A third biotope containing maerl gravel was recorded at the south end of Holy Isle and on the Isle’s north-east coast (SS.SMp.KSwSS.LsacR.Gv - Laminaria saccharina and robust red algae on infralittoral gravel and pebbles). In these semi-exposed areas the maerl gravels possessed significant over-growths of the invasive algal species Heterosiphonia japonica and Trailliella intricata.

Relatively rich infaunal communities, dominated by large bivalves were found to be present within the underlying sediments at all dive sites.

A seagrass bed in Whiting Bay was surveyed opportunistically as an alternative to the remoter and exposed maerl gravel areas around the Iron Rock Ledges. Survey logistics and weather prevented access to these. A healthy, rich epifaunal and floral community was found to be associated with the Zostera marina bed in Whiting Bay.

For further information on this project contact: Lisa Kamphausen, Scottish Natural Heritage, Great Glen House, Leachkin Road, Inverness, IV3 8NW.

Tel: 01463 725014 or [email protected] For further information on the SNH Research & Technical Support Programme contact:

Research Coordinator, Scottish Natural Heritage, Great Glen House, Leachkin Road, Inverness, IV3 8NW. Tel: 01463 725000 or [email protected]

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Table of Contents Page

1. INTRODUCTION 1

2. METHODS 3 2.1 MNCR phase II – style diver transect surveys 4 2.2 Diver transect quantitative maerl assessments 4 2.3 Maerl biotope infaunal diver coring and epifloral sampling and video

records 5 2.4 Whiting Bay Zostera bed - method variations 5 2.5 Field laboratory methods 5 2.6 Laboratory methods 6

3. RESULTS 8 3.1 Physical characteristics of maerl habitats 8 3.2 Phase II dive survey epifaunal and floral abundance data 8 3.2.1 Pladda (station PL34) 10 3.2.2 Holy Isle South (station HI22) 10 3.2.3 Holy Isle North (stations HIN04 and HIN02) 11 3.2.4 Lamlash Bay North (station LBN) 12 3.2.5 Microscopic epiphytic maerl algae 13 3.2.6 Whiting Bay seagrass bed (station WBZ) 14 3.3 Particle size distribution 15 3.4 Macrobenthic infauna 16 3.4.1 Univariate analysis of core data 16 3.4.2 Multivariate analysis of the core data 16

4. DISCUSSION 20 4.1 Study of maerl beds in the South Arran MPA 20 4.2 Comparison of the ‘maerl bed’ and ‘maerl gravel’ biotopes 21 4.3 Macrobenthic infauna - comparison to other Arran and UK maerl bed

records 22 4.4 Whiting Bay seagrass bed 23

5. REFERENCES 24

ANNEX 1: RAW DATA 26

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1. INTRODUCTION

The South Arran Marine Protected Area (MPA) (Figure 1) is one of three MPAs within the Clyde Sea, together with the Clyde Sea Sill MPA to the south and Upper Loch Fyne and Loch Goil MPA to the north.

Figure 1. The South Arran MPA.

The ‘maerl beds’ within the MPA are largely considered to be in poor condition and a ‘recover’ conservation objective has been assigned to this feature by SNH. Site management proposals consulted upon at the end of 2014 (Marine Scotland, 2014), included known maerl beds and larger ‘recovery areas’ located around clusters of records of sparse living maerl and relatively unbroken dead maerl which are deemed to have the highest potential to aid recovery. These maerl habitats were initially afforded protection through an urgent Marine Conservation Order1 which took effect on 1 October, 2014 after a voluntary fisheries management agreement was breached. The habitats are now protected under the fisheries management measures which came into force in December 2015 (Scottish Statutory Instruments, 2015). Survey work undertaken in connection with the Scottish MPA Project since 2012 greatly improved understanding of the distribution of all seabed habitat features within the MPA, but records were still concentrated on the existing Lamlash Bay No Take Zone (NTZ). To better define the full distribution and extent of relevant seabed habitats throughout the MPA and to inform future site management decisions, a broad coverage drop-down video survey was conducted in 2014 (reported separately as Morris-Webb, 2015). That study, which was undertaken just prior to the more detailed diving studies reported upon here, helped to target suitable locations for diver sampling.

1 See http://www.gov.scot/Topics/marine/marine-environment/mpanetwork/southarranmco

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The principal aim of the presented study was to assess the levels of biodiversity associated with maerl and maerl gravel biotopes around Arran in order to create a baseline against which the effectiveness of future management measures can be assessed. The collection of detailed diver observations from the subtidal Zostera marina seagrass beds within Whiting Bay, the largest known example of this feature in the Clyde Sea, was a contingency objective for adverse weather conditions, to provide context to extent mapping of this habitat undertaken earlier (Morris-Webb, 2015). Aquatic Survey and Monitoring Limited were commissioned to conduct detailed assessments of various sites representative of the respective protected features, using in situ diving observations and collection of core samples for infaunal analyses. In the field season of 2015 SNH collected additional drop video, grab and diving data to establish a pre-management baseline of maerl bed, maerl gravel, and burrowed mud feature condition in the South Arran MPA. Results of these surveys will be reported on separately once analysis is concluded, and all pre-management data will then be combined to establish a baseline.

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2. METHODS

Between September 21 - 26, 2014 MNCR phase II style dive surveys were conducted at six locations within the South Arran MPA (Table 1 and Table 2, Figure 2). Survey stations were selected to provide representation of a range of environmental conditions and pressure regimes, inside and outside areas where existing and potential future protection measures were expected to be in place, while making concessions to accessibility in the prevailing weather. The final management measures will now afford protection from bottom contacting mobile fishing to all maerl bed and maerl gravel areas in the MPA, but drop video and grab work in 2015 also contains stations outside the MPA that act as un-managed comparison sites. Precise locations of stations in this survey were based on data collected prior to the dive survey by an SNH drop-down video survey (Morris-Webb, 2015). That work identified new high quality examples of maerl beds in the MPA which had not been surveyed in detail previously and hence were visited during this dive survey.

Table 1. Location of dive sites targeted during the 2014 Arran survey.

Dive site name Date Site code Latitude Longitude Depth BCD (m) Pladda 21/09/2014 PL34 55o 26.181’ -5o 7.496’ 7.5 Holy Isle South 22/09/2014 HI22 55o 30.646’ -5o 3.727’ 7.8 Holy Isle North (inside the NTZ*) 23/09/2014 HIN04 55o 32.320’ -5o 4.655’ 6.2 Lamlash Bay North maerl bed 24/09/2014 LBN 55o 32.760’ -5o 5.314’ 9.8 Holy Isle North (outside the NTZ*) 25/09/2014 HIN02 55o 32.138’ -5o 4.339’ 6.0 Whiting Bay Zostera bed 26/09/2014 WBZ 55o 29.852’ -5o 5.273’ 4.4

* No Take Zone A further two reconnaissance dives were undertaken to determine the eastern and western extent of the Lamlash Bay North (LBN) maerl bed. Positional information for these two diving stations is provided below (Table 2). A pair of divers swam along the maerl bed following the 8 and 12 m contours respectively until live maerl coverage dropped below 5%. At that point they moved to the middle of the maerl band at about 10 m depth, signalled the end of the bed with tugs on their SMB, and the position was recorded by a snorkeler carrying a GPS.

Table 2. 2014 dive sites marking the eastern and western margins of the Lamlash Bay North maerl bed (LBN).

Dive site name Date Site code Latitude Longitude Depth BCD (m) (at middle of maerl band)

Lamlash Bay North maerl bed

23/09/2014 Eastern edge

55o 32.750’ -5o 5.192’ 9.8

Lamlash Bay North maerl bed

25/09/2014 Western Edge

55o 32.768’ -5o 5.428’ 9.8

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Figure 2. Dive survey positions presented with relevant biodiversity data.

2.1 MNCR phase II – style diver transect surveys

The survey was conducted from the 5.8 m rib, North Shore. Survey sites were identified with the boat’s chart plotter (Standard Horizon CP180i GPS), and marked with a buoyed shot to which a 25 m tape measure and leaded safety line were attached. Surveys were conducted along fixed 25 m transects. A pair of divers descended the shot line and selected a bearing for the 25 m transect which was most representative of the target survey feature. This depended on the topography of the seabed and features such as mats or clumps of algae such as Saccharina latissima which were avoided. The tape and leaded safety line were then laid out along the selected bearing. A small weight was attached to the end of the tape and line to keep them in place. The divers then returned to the origin of the transect and began the Phase II survey. During the survey, divers clipped themselves to the leaded line with buddy lines to be in contact with the surface via the shot line if necessary. The 25 m transect was surveyed for a width of 2 m either side of the tape. The abundance of all taxa observed was recorded according to the MNCR Abundance Scale (SACFOR). The divers, an algologist and a zoologist, surveyed the seabed for flora and fauna on one side of the tape each, and upon reaching the end of the tape swapped sides and continued the survey along the opposite side of the transect. The pair of divers was equipped with a digital camera and flash, with which they took a photographic record of much of the faunal and floral component of the biotope. 2.2 Diver transect quantitative maerl assessments

When the first pair of divers returned to the boat, a second wave of three divers entered the water. Two of the divers took with them a 0.25 m2 quadrat and camera each. They recorded percentage cover of live and dead maerl in 20 quadrats placed at pre-selected random

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distances along the transect, and took a picture of each quadrat. The pictures were reviewed at a later time as a quality control measure of the maerl abundance assessments made in the field. 2.3 Maerl biotope infaunal diver coring and epifloral sampling and video records

The third diver of the second wave collected infaunal sediment cores, a sediment sample for particle size distribution (PSD) analysis, and algal samples. The 10 cm diameter corers were driven into the sediment to a depth of 20 cm (minimum) before they were securely capped, extracted from the sediment and retained. The PSD sample consisted of a sediment sample taken from the top 5 cm of the undisturbed seabed. The sediment core samples and PSD samples were all collected from random locations within 5 m of the shot weight. Some of the small and finely branched seaweeds associated with maerl can only be identified with a microscope. To study these, samples of maerl and sediment were collected from five randomly selected decimetre squares in the vicinity of the transect. The surface 30 mm of maerl within these decimetre squares were sampled carefully by hand and placed in mesh bags with a 1 mm mesh size. The core samples, PSD sample and maerl algae bags were carefully returned to the surface with the aid of a diver lifting bag. On finishing the infaunal macrobenthic and maerl sampling, a video record was made of both sides of the transect with a handheld HD video camera. 2.4 Whiting Bay Zostera bed - method variations

To select a location for a survey of the Zostera bed location in Whiting Bay, the presence of a suitable dense Zostera marina sward was confirmed using both a bathyscope over the side of the RIB, and with snorkelers in the water. Once on location, a buoyed shot was dropped in the sand between patches of seagrass to mark the dive site. Because of the small-scale patchiness of the seagrass, a fixed 25 m transect was not suitable to record details of the seagrass bed. Therefore the Phase II information was collected by the first pair of divers from within a 50 m radius of the shot marker, but only within the seagrass bed patches. A second pair of divers collected Zostera marina shoot density records within 46 0.25 m2 quadrats. The quadrats were haphazardly thrown into seagrass patches within the survey area. A video record was also made of the Zostera biotope. 2.5 Field laboratory methods

Following the daily fieldwork, samples were returned to the field laboratory and variously processed: Specimens collected during the dives for taxonomic verification were identified using

both high and low powered microscopes and the latest taxonomic keys. Field data sheets and notes were updated and data were entered into Excel spreadsheets.

Photographs were downloaded, renamed and catalogued in an Excel spreadsheet. The photos were processed using a set protocol. Jpg photograph files were renamed with the following convention: 20140925_FDB_HIN02_0074.jpg = 8 digit date (YYYYMMDD), 3 letter initials of photographer (Francis Daly Bunker), Site code (Holy

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Isle North 02) and a 3 or 4 digit consecutive photo ID number derived from the camera (0074) followed by the file extension( .jpg). No spaces were allowed in the file name. The photographs were catalogued in the relevant Excel spreadsheet that included a hyperlink that allowed the photographs to be opened directly from the Excel file when stored in the appropriate sub-directory.

Maerl algae samples. The maerl algae bags had a 1 mm mesh and so the fine particles of sediment were removed by gentle agitation in seawater. The remaining larger particles (mainly fragments of shell, gravel and maerl) were examined under a stereomicroscope and species lists of the seaweeds present were compiled. Any macrofauna present were also noted for completeness.

Macrobenthic infaunal cores were extracted and sieved over a 1 mm mesh. The sieved residue was placed in a container labelled inside and out with the site name, replicate number, date taken and sieve mesh size. The samples were then fixed in a 10% formaldehyde solution and labelled with both toxic and hazardous labels.

2.6 Laboratory methods

Macrobenthic taxonomic analysis. The analysis of marine macrobenthic samples adhered to the mandatory sections of the NMBAQC Processing Requirement Protocol (Version 1.0) which conforms to EN ISO 16665:2005 and BS EN 14996:2006. All five replicate cores taken at each location were processed and analysed. Each of the fixed and preserved sediment samples was carefully washed in freshwater to remove the formalin, transferred into labelled buckets and Rose Bengal was added to stain the biological material in order to facilitate the sorting process. Samples were then placed in gridded white trays, systematically examined by eye and all fauna picked out, sorted into phyletic groups and placed in vials in 70% industrial methylated spirit. The extracted biota were identified to the lowest possible taxonomic group using a combination of binocular stereo microscopes and high magnification binocular compound microscopes and counted. Where possible all organisms were identified to species level according to the nomenclature of the World Register of Marine Species (WoRMS) with species codes (where available) consistent with Howson & Picton (1997). Taxonomic literature used was consistent with the NMBQAC Taxonomic Literature database (version 107), including use of relevant journal papers, unpublished workshop keys (primarily from NMBAQC workshops), internet keys and internal documents compiled from these sources. All whole organisms were identified and counted. Damaged or incomplete specimens were identified and counted only if the head was present; otherwise parts of organisms were recorded but not enumerated. Juveniles were recorded as such when identification to species level was not possible due to the under-development of key features, or if the specimen was less than 10% of the average adult body size.

Particle size distribution analysis was carried out by Hebog Environmental Ltd. using a dry sieving technique and sieves conforming to the Wentworth scale. Approximately 80 g of sediment was oven dried at 100 °C until constant weight was reached. The dried sediment was weighed and washed through a 63 µm sieve. The effluent, which contained the mud/silt fraction (< 63 µm) was not retained. The remaining sediment was again oven dried at 100 °C until constant weight was reached. This weight was recorded and the percentage of the mud/silt fraction (< 63 µm) was calculated as the difference between these two weights. The fractions > 63 µm were transferred to the coarsest of a series of stacked sieves, placed on an automatic shaker for 15 minutes and the contents of each sieve weighed. Fractions are > 8 mm, 4 - 8 mm, 2 - 4 mm,

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1000 - 2000 µm, 500 - 1000 µm, 250 - 500 µm, 125 - 250 µm, 63 - 125 µm and < 63 µm. All fractions were weighed to an accuracy of ± 1 mg.

Loss on ignition analysis was also carried out in order to determine the amount of organic matter found in the sediment and available to the macrobenthic infauna. Approximately 10 g of sediment was placed into a ceramic crucible and dried at 100°C in an oven. The sample was weighed to an accuracy of ± 1 mg and heated to 550°C for four hours. The crucible was placed over a desiccant to cool and then re-weighed (methodology is based on Holme and McIntyre (1984) Methods for the Study of Marine Benthos, Chapter 3: Sediment Analysis. IBP Handbook 16, Blackwell Scientific Publications, p. 41-65).

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3. RESULTS

All raw data are presented in tabular form in Annex 1. 3.1 Physical characteristics of maerl habitats

Table A in Annex 1 presents the site characteristics data for all five of the ‘maerl’ and ‘maerl or coarse shell gravel with burrowing sea cucumbers’ sites. The groups fall into three distinct sediment categories:

1. Pladda (PL34): this site was a tide swept, exposed and level seabed consisting mostly of maerl gravel and live medallions of Phymatolithon calcareum maerl (quadrats determined 29 % live maerl, and 92 % combined live maerl and maerl gravel), with some pebbles and a substantial amount of lose empty shells covering the maerl gravel (30 %).

2. Holy Isle (HI22, HIN04 and HIN02): these sites all had less whole shell on the surface (~ 20 %) and were made up of close to 100 % maerl gravel, but contained very little live Phymatolithon calcareum maerl (< 1 – 2 %). Fragments of Lithothamnion glaciale were also seen at HIN04. All three sites were determined to be part of the ‘kelp and seaweed communities on sublittoral sediment’ protected feature.

3. Lamlash Bay (LBN): this site had a very high live Lithothamnion glaciale2 maerl content (55 – 75 %) and lower maerl gravel content (~25 %), a slightly lower whole shell component and slightly higher mud component than the other maerl sites, a fact borne out by the silt/clay fraction result for this site (see Annex 1 Table E - diver core PSD). The increased softness at this site was highly noticeable during the dive. The bed sloped from north to south into the channel between Arran and Holy Isle.

3.2 Phase II dive survey epifaunal and floral abundance data

The Phase II survey produced records of 186 taxa within the five maerl and maerl gravel sites dived. Taxa abundance was collected as SACFOR values (JNCC, 1990), hence multivariate statistics are not appropriate for analysis. The full set of results is shown in Annex 1 Table C. Table 3 below contains the abundance values of the dominant flora and fauna at each site. Only those recorded at Super abundant (S) to Frequent (F) are presented. The following sections describe the flora and fauna at each station in further detail.

2 Confirmed by ‘thin section analysis’ after acid digestion by F. Bunker, J Hall-Spencer and Dr

Viviana Peña.

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Table 3. SACFOR abundances of dominant flora and fauna recorded by Phase II style dive surveys at each of the South Arran dive sites in 2014. Maerl entries are pink, other algae green, and animals white. S = super abundant, A = abundant, C = common, F = frequent, O = occasional, R = rare.

PL34 LBN Phymatolithon calcareum C Lithothamnion glaciale S

Marthasterias glacialis C Bonnemaisonia hamifera (Trailiella intricata)

S

Pomatoschistus sp. C Chaetopterus variopedatus C

Heterosiphonia japonica F Galathea intermedia C

Saccharina latissima F Hydroides norvegicus C

Scinaia interrupta F Inachus dorsettensis C

Cerianthus lloydii F Necora puber C

Chaetopterus variopedatus F Spirobranchus sp. C

Necora puber F Spirorbinae C

Spirobranchus sp. F Saccharina latissima F

Testudinalia testudinalis F Pomatoschistus sp. F

Tectura virginea F

HI22 HIN04 HIN02

Phymatolithon calcareum R Lithothamnion glaciale R Phymatolithon calcareum O

Cerianthus lloydii A Phymatolithon calcareum R Heterosiphonia japonica C

Saccharina latissima C Heterosiphonia japonica S Saccharina latissima C

Desmarestia aculeata F Saccharina latissima C Ascidiella aspersa C

Heterosiphonia japonica F Asterias rubens C Chaetopterus variopedatus C

Ascidiella aspersa F Cancer pagurus F Tectura virginea C

Chaetopterus variopedatus F Cerianthus lloydii F Cancer pagurus F

Dosinia exoleta F Luidia ciliaris F Marthasterias glacialis F

Gibbula magus F Marthasterias glacialis F Necora puber F

Necora puber F Necora puber F Pecten maximus F

Polititapes rhomboides F Pomatoschistus sp. F Pecten maximus (juv) F

Pomatoschistus sp. F Porania pulvillus F Pomatoschistus sp. F

Spirobranchus sp. F Spirobranchus sp. F

WBZ Zostera marina A

Echinocardium cordatum C

Echiurus echiurus C

Ensis arcuatus C

Liocarcinus depurator C

Macropodia rostrata C

Pagurus bernhardus C

Pomatoschistus sp. C

Rissoa lilacina C

Rissoa parva C

Heterosiphonia japonica F

Saccharina latissima F

Necora puber F

Ophiura ophiura F

Pholis gunnellus F

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3.2.1 Pladda (station PL34)

SS.SMp.Mrl.Pcal.R This station was a shallow (circa 8 m BCD) tide-swept maerl bed, with substantial growths of the kelps Saccharina latissima and Saccorhiza polyschides, as well as the brown alga Desmarestia aculeata on cobbles and boulders, along with encrusting corallines. The combined live maerl and maerl gravel component of the substrate made up 92 %, and live maerl coverage was 29 %, i.e. SACFOR Common (avg. of 20 quadrats – see Annex 1 Table B), in the form of Phymatolithon calcareum medallions on the sediment surface, along with both Ulva sp. and Scinaia interrupta which were frequent on the pebbles and amongst the sands and gravels. The invasive non-native red alga Heterosiphonia japonica was also found at a SACFOR abundance of Frequent on the shells, pebbles and cobbles. The fauna was dominated by the large mobile starfish Marthasterias glacialis and small mobile Pomatoschistus gobies. Decapods such as Cancer pagurus, Necora puber and Paguridae roamed over the sediment, whilst cobbles were colonised by Testudinalia testudinalis, Gibbula magus, Spirobranchus sp., Ascidiella aspersa and Celleporella hyalina. Within the sediment surface, tubes of Chaetopterus variopedatus protruded with and the anemones Sagartiogeton spp. and Cerianthus lloydii were found. Figure 3 shows representative images from the Pladda site.

Figure 3. Illustrative shots of Pladda maerl bed (PL34) showing medallions of Phymatolithon calcareum and empty shells, and Scinaia interrupta in the left image.

3.2.2 Holy Isle South (station HI22)

SS.SMp.KSwSS.LsacR.Gv This was another shallow station was at about 8 m BCD, and differed from the station at Pladda in its algal complement only by the lack of the large kelp fronds of Saccorhiza polyschides and the amount of live maerl present. Live maerl was sparse at this site, being recorded as rare (R) and 2 % (avg. of 20 quadrats, see Annex 1 Table B) compared to Common (C) with 29 % (avg) at Pladda. The combined live maerl and maerl gravel component of the substrate made up 97 % of the substrate here, similar to the 92 % at Pladda. The fauna differed from that at Pladda in that there were several large bivalve species, notably Dosinia exoleta, Polititapes rhomboides and Pecten maximus. These were often present at the surface of the sediment. This in turn has led to increased numbers of the starfish Asterias rubens and Marthasterias glacialis in the biotope. These differences aside, the biotopes were very similar. Figure 4 shows representative images from the Holy Isle South site.

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Figure 4. Illustrative shots of the Holy Isle South (HI22) site, showing maerl gravel with scattered shells, and a Luidia luidia star fish on the left.

3.2.3 Holy Isle North (stations HIN04 and HIN02)

SS.SMp.KSwSS.LsacR.Gv Physically similar to the Holy Isle South station, the stations north of Holy Isle, HIN04 inside the No-Take Zone (NTZ) and HIN02 outside the NTZ, were of the same biotope with some differences in dominant biota. 86 and 85 taxa respectively were recorded on the transects, and Table 3 shows that the conspicuous dominant taxa were very similar between the sites. In terms of the algal component, the main difference between the sites was the amount of Heterosiphonia japonica present at each site; Super-Abundant at HIN04, whilst only Common at HIN02. Both these sites contained very low quantities of live Phymatolithon calcareum maerl (< 3 %), and were dominated by encrusting corallines and Saccharina latissima. Small amounts of Lithothamnion glaciale maerl (< 1 %) were also recorded at HIN04. The presence of Lithothamnion glaciale has been confirmed by thin section analysis of the maerl. In faunal terms the main differences between HIN04 and HIN02 were the presence and absence of several large bivalves such as Pecten maximus or Polititapes rhomboides. Both Aequipecten opercularis and Pecten maximus were recorded at HIN02, outside the NTZ and were not recorded at HIN04, inside the NTZ on this occasion. The biotope fauna was dominated by large echinoderms (Marthasterias glacialis, Asterias rubens, Luidia ciliaris and Porania pulvillus) and bivalves (Pecten maximus, with Polititapes rhomboides, and Ensis sp. also present) as well as mobile decapod crustaceans (Necora puber and Cancer pagurus). Chaetopterus variopedatus and sedentary ascidians (Ascidiella aspersa and Corella parallelogramma) punctuated the sediment surface. Individuals of Limaria hians were found at both stations, though no flame shell beds were found in the vicinity during a subsequent search by two of the divers. Figure 5 shows images representative of the Holy Isle North sites.

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Figure 5. Illustrative shots of the northern Holy Isle sites HIN02 (top row) and HIN04 (bottom row), showing maerl gravel and associated species (Scinaia interrupta top row and Luidia luidia bottom row).

3.2.4 Lamlash Bay North (station LBN)

SS.SMp.Mrl.Lgla This was by far the densest maerl bed surveyed in South Arran. It was dominated by live Lithothamnion glaciale maerl with cover of between 55 - 75 %. There was, however, also a considerable over-growth of Bonnemaisonia hamifera (Trailliella intricata) red alga across the bed, which is of concern. Other dominant algae included Saccharina latissima, Cutleria multifida, Corallina officinalis and Heterosiphonia japonica. The fauna was very similar to that found in the other maerl beds with large echinoderms, such as Asterias rubens and Marthasterias glacialis. Astropecten irregularis was also present and recorded only at this station during the survey. Several decapod crustaceans not recorded at other stations (Galathea intermedia, Liocarcinus depurator, Pandalus montagui and Anapagurus hyndmanni) were seen. Sedentary fauna included the tube worms Chaetopterus variopedatus, Spirobranchus spp., spirorbinae and Hydroides norvegicus and ascidians such as Ascidiella aspersa. Figure 6 shows images representative of the Lamlash Bay north site. This dense maerl bed had not previously been known to SNH but was shown to the survey team by local conservationists. Since it contains substantially more live maerl than any other bed currently known in the Clyde the survey team delineated its extent by conducting dives along it, and marking the position where live maerl cover dropped below 5 %. They found the bed to be a well-defined band between roughly the 7 and 12 m depth contour BCD of the steeply sloping seabed, stretching for a total of 250 m from NW to SE (Figure 7).

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Figure 6. Illustrative shots of the Lamlash Bay North maerl bed (LBN), showing dense live maerl thalli, scattered shells, and a cushion star (Porania pulvillus).

Figure 7. Details of the Lamlash Bay North (LBN) maerl bed.

3.2.5 Microscopic epiphytic maerl algae

Tables 4 and 5 present the summary results of the maerl algae sample analysis (5 x 10 cm2

per site). The species list is presented in Annex 1 Table D. Forty-seven algal species / taxa were recorded and eleven faunal taxa. Table 4 shows numbers per site. Nineteen of the microscopic algal taxa were not recorded by the diving Phase II survey. These new taxa are presented in Table 5 below, and include the endemic maerl epiphyte Gelidiella calcicola. These data add to the knowledge base of the maerl bed quality within the South Arran MPA.

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Table 4. Number of epiphytic algal and faunal taxa recorded on the maerl thalli.

PL34 HI22 HI04 All taxa 12 12 16 18 13 11 17 14 10 15 25 20 12 19 17 Animal taxa 2 1 1 1 5 Algal taxa 10 11 15 18 13 11 17 13 10 15 20 20 12 19 17 Mean 14.2 13.4 18.6

Table 4 continued.

LBN HI02 All taxa 14 13 15 12 14 18 12 17 14 12 Animal taxa 1 2 2 3 Algal taxa 14 13 14 12 14 16 12 15 11 12 Mean 13.6 14.6

Table 5. Epiphytic algae found on maerl / maerl gravel sediments.

Additional algal species recorded only by microscopic analysis of maerl samples

Acrosorium venulosum Falkenbergia Ptilota gunneri Aglaothamnion byssoides Gelidiella calcicola Sphacelaria plumosa Aglaozonia sp. Griffithsia corallinoides Trailliella intricata Audouinella sp. Halarachnion ligulatum Ulva lactuca Cladophora sp. Plocamium cartilagineum Ulva sp. (Tubular) Cystoclonium purpureum Polysiphonia harveyi Ectocarpaceae Ptilothamnion sphaericum

3.2.6 Whiting Bay seagrass bed (station WBZ)

SS.SMp.SSgr.Zmar The seagrass bed was found in a shallow sandy bay at a depth of ~ 4.5 m (BCD). Table 3 presents the SACFOR abundances of the dominant taxa recorded during the Phase II survey and the full list is presented in Annex 1 Table C. The bed is made up of a mosaic of patches of dense seagrass fronds interspersed by smaller areas of sand. The Zostera marina shoot density within the seagrass patches was 89 shoots m-2 at the survey site at north end of Whiting Bay. In terms of algae, the biotope was dominated by Heterosiphonia japonica and Saccharina latissima, with numerous small red, brown and green algal species occurring at low abundances. Twenty-two algal taxa were recorded during the survey. Thirty seven faunal taxa were recorded by the divers and this community was dominated by large mobile crustaceans such as Pagurus bernhardus, Macropodia rostrata, Liocarcinus depurator, Carcinus maenas and Necora puber. The infaunal community of the sand contained several megafaunal species, of which the most notable were dense aggregations of Echinocardium cordatum and what was thought to be the echiuroid Echiurus echiurus, though no live specimens were obtained for confirmation. Figure 8 shows the typical proboscis marks in the sand.

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Figure 8. Proboscis marks typical of Echiurus echiurus in the Whiting Bay seagrass bed.

Bivalves such as Ensis arcuatus were also noted in the sediment, as were the tubes of the polychaete Chaetopterus variopedatus. The leaves of the Zostera marina plants were frequently covered with dense aggregations of Rissoa lilacina and Rissoa parva, whilst the surface of the sand was dominated by Ophiura ophiura, Asterias rubens, tubed anemones, several other starfish and numerous small fish (e.g. Pomatoschistus sp., Pholis gunnellus). Occasional pebbles and cobbles were colonised by barnacles (Balanus crenatus), bryozoans (Celleporella hyalina), ascidians (Ascidiella aspersa), and tubed polychaetes (Spirobranchus sp.).

Figure 9. Illustrative shots of the Zostera marina bed in Whiting Bay.

3.3 Particle size distribution

The results of particle size distribution and loss on ignition analyses (assessing organic matter) of the infaunal cores are presented in Annex 1 Table F. The particle size distributions confirmed the divers’ in situ observations that the survey sites lie along a gradient of exposed, semi-exposed and sheltered sites. Cores from Pladda (PL34), the exposed site, were made up of coarse sediments, dominated by pebbles, gravel and very coarse sand. Cores from the semi-exposed Holy Isle sites (HI22, HIN04 and HIN02) contained fine gravels and coarse sands with slightly greater silt and clay components as well as an increased organic matter fraction. Cores from the sheltered

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Lamlash Bay North (LBN) site still contained a fine gravel / coarse sand fraction (due to the maerl component), but also had far greater amounts of silt and clay ( ~ 16 % ) and organic matter ( ~ 3 % ). 3.4 Macrobenthic infauna

Analysis of the >1 mm macrobenthic infauna revealed a diverse community of invertebrates within the maerl and maerl gravel sediments. The table of taxa and their abundances for each core is shown in Annex 1 Table G. In total 130 taxa were identified in the 25 cores. The data were analysed with the marine invertebrate community analytical computer package PRIMER 6. 3.4.1 Univariate analysis of core data

To calculate indices of community diversity, a univariate analysis (the Diverse protocol of PRIMER 6) was conducted on all individual cores, and at site mean values are shown in Table 6. Values for all indices were highest for either the Lamlash Bay (LBN) cores or for the Pladda (PL34) cores, and lowest for Holy Isle sites. Pielou’s evenness statistic showed that station HIN02 was the least even, i.e. most dominated community, with nematodes, the spionid Aonides paucibranchiata and the brittlestar Amphipholis squamata being the most likely candidate taxa for this effect. This pattern is supported by the Shannon-Wiener and the Simpson’s diversity indices, which yield the highest diversity scores for LBN and the lowest for HIN02.

Table 6. Mean univariate characteristics of the site core data.

Stn. S N d J' H'(loge) 1-Lambda' S Tot

PL34 31 (26-34) 106 (83-

142) 5.87 0.86 2.83 0.92 52

HI22 25 (21-28) 56 (35-92) 5.66 0.91 2.78 0.93 50

LBN 30 (24-35) 61 (45-77) 6.78 0.93 3.09 0.96 71

HIN02 26 (22-32) 104 (46-

183) 5.27 0.76 2.43 0.85 58

HIN04 32 (25-38) 94 (73-105) 6.07 0.81 2.70 0.89 60 S - Mean total taxa: taxa with non zero counts. (Range shown in brackets) N - Mean total individuals: The mean number of individuals per core. (Range shown in brackets). d - Mean Margalef's richness for each sample. (S-1)/Log(N) - it is a measure of the number of taxa present, making some allowance for the number of individuals. J’ - Mean Pielou's evenness - this is a measure of equitability, a measure of how evenly the individuals are distributed among the different taxa. H'(loge) - Mean Shannon-Wiener diversity index 1-Lambda'- Mean Simpson's diversity index S Tot - Total number of taxa recorded in all five cores per site

3.4.2 Multivariate analysis of the core data

Further trends in the infaunal community were illustrated using multivariate analysis of the data. A Bray Curtis similarity analysis was performed to produce a similarity matrix. This was run through a group average clustering routine in PRIMER 6 to produce a dendrogram illustrating groupings in the data (Figure 10). As expected from the epifaunal and floral characteristics, the Pladda site and the Lamlash Bay North site separate out from the Holy Isle sites. The infaunal communities they support clearly both have a greater intra-group similarity than inter-group similarity. The Holy Isle

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South site HI22 also separates out, and although the two northern sites HIN02 and HIN04 are generally very similar, there are a few outliers, such as HIN04#4, HIN02#1 and HIN02#5.

South Arran - 'diver cores' macrobenthic dataGroup average clustering

LBN

#4

LBN

#1

LBN

#3

LBN

#2

LBN

#5

Pl3

4#1

Pl3

4#3

Pl3

4#5

Pl3

4#2

Pl3

4#4

HI0

4#4

HI0

2#1

HI0

2#2

HI0

2#3

HI0

2#4

HI0

4#1

HI0

4#3

HI0

4#2

HI0

4#5

HI0

2#5

HI2

2#2

HI2

2#4

HI2

2#3

HI2

2#1

HI2

2#5

Samples

100

80

60

40

20

Sim

ilarit

y

Transform: Square rootResemblance: S17 Bray Curtis similarity

Figure 10. Infaunal cores macrobenthic data – group average clustering Bray Curtis similarity.

South Arran 'diver cores' macrobenthic data2 dimensional MDS plot

Transform: Square rootResemblance: S17 Bray Curtis similarity

clustersBACDEGF

Pl34#1

Pl34#2Pl34#3

Pl34#4Pl34#5

HI22#1HI22#2

HI22#3

HI22#4HI22#5

LBN#1

LBN#2

LBN#3

LBN#4

LBN#5

HI02#1

HI02#2HI02#3HI02#4

HI02#5

HI04#1HI04#2

HI04#3

HI04#4

HI04#5

2D Stress: 0.19

Figure 11. Infaunal cores macrobenthic data - two dimensional MDS plot.

A multidimensional scaling (MDS) analysis (Figure 11), a means of visualising similarity across samples, produces similar groupings as the dendrogram, where the cores from Pladda and Lamlash Bay separate out from a looser cluster of Holy Isle cores.

A B C F G D E Cluster group

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To determine the taxa responsible for the clustering patterns, the Bray Curtis similarities were subjected to a SIMPER analysis within PRIMER 6. The results are shown in the following Tables 7 - 10. Only taxa with a similarity percentage contribution of over 5%, or making up the top cumulative 50%, are included. Clusters with fewer than two samples could not be analysed through SIMPER, but their dissimilarities from the nearest clusters were determined. The dendrogram (Figure 10) illustrates that the samples LBN#5, HIN02#1, HIN04#4 (A, F and G) are outliers, and Tables 11 - 13 present the taxa that create the dissimilarities from their nearest clusters. This analysis shows that the exclusion of LBN#5 from Group B, and the exclusion of HIN02#1 and HIN04#4 from Group D, are caused by the presence or absence of a few otherwise common taxa, which reflects the patchy nature of species distribution in the marine environment.

Table 7. SIMPER taxa for Group B clustered cores.

Group B - Lamlash Bay North Average similarity: 47.84 Species Av.Abund Av.Sim Sim/SD Contrib% Cum.% Galathea intermedia 2.27 5.65 7.37 11.81 11.81 Ophiothrix fragilis 2.39 5.2 7.2 10.88 22.69 Amphipholis squamata 1.73 3.67 4.22 7.67 30.37 Lysianassa plumosa 1.6 3.43 4.19 7.16 37.53 Nematoda 1.29 2.92 5.5 6.11 43.64 Socarnes erythrophthalmus 1.64 2.92 5.5 6.11 49.74 Clausinella fasciata 1.1 2.73 10.79 5.71 55.46 Dosinia exoleta 1.43 2.52 0.91 5.27 60.73

Table 8. SIMPER taxa for Group C clustered cores.

Group C - Pladda PL34 Average similarity: 55.29 Species Av.Abund Av.Sim Sim/SD Contrib% Cum.% Nematoda 4.03 7.84 4.95 14.18 14.18 Socarnes erythrophthalmus 2.83 5.29 3.65 9.57 23.74 Animoceradocus semiserratus 2.07 3.91 3.44 7.08 30.82 Dosinia exoleta 2.1 3.79 3.02 6.85 37.67 Pisione remota 2.09 3.44 2.75 6.23 43.9 Polygordius 2.01 3.03 2.21 5.48 49.39 Clausinella fasciata 1.33 2.37 7.33 4.28 53.67

Table 9. SIMPER taxa for Group D clustered cores.

Group D - HI04 and HI02 #2#3#4 Average similarity: 53.79 Species Av.Abund Av.Sim Sim/SD Contrib% Cum.% Nematoda 5.34 12.1 10.87 22.49 22.49 Amphipholis squamata 2.83 5.9 3.23 10.98 33.47 Aonides paucibranchiata 2.22 4.8 6.28 8.92 42.39 Sphaerosyllis bulbosa 1.98 4.16 5.13 7.73 50.12 Dosinia exoleta 2.04 3.89 2.3 7.24 57.36

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Table 10. SIMPER taxa for Group E clustered cores.

Group E - HI22 and HI02 #5 Average similarity: 41.96 Species Av.Abund Av.Sim Sim/SD Contrib% Cum.% Nematoda 1.69 5.02 4.07 11.97 11.97 Mediomastus fragilis 1.26 4.06 4.59 9.68 21.65 Amphipholis squamata 1.92 3.28 1.08 7.81 29.46 Thracia villosiuscula 1.3 3.24 1.28 7.73 37.19 Polygordius sp. 1.33 3.22 1.3 7.67 44.86 Clausinella fasciata 1.24 3.05 1.27 7.26 52.12 Lysidice unicornis 1.02 2.62 1.34 6.24 58.35 Aonides paucibranchiata 1.07 2.49 1.36 5.93 64.28

Table 11. The taxa producing the top 20% of the dissimilarity between A & B.

Groups B & A Average dissimilarity = 63.17 LBN#5

Group B Group A Species Av.Abund Av.Abund Av.Diss Diss/SD Contrib% Cum.% Galathea intermedia 2.27 0 2.99 8.35 4.73 4.73 Podarkeopsis helgolandicus 0 1.73 2.3 12.34 3.64 8.37 Socarnes erythrophthalmus 1.64 0 2.16 1.65 3.43 11.79 Leptochiton cancellatus 0 1.41 1.88 12.34 2.97 14.76 Thracia villosiuscula 0 1.41 1.88 12.34 2.97 17.73 Spirobranchus lamarcki 1.34 0 1.76 1.15 2.79 20.52

Table 12 The taxa producing the top 20% of the dissimilarity between F & D.

Groups D & F Average dissimilarity = 56.51 HI04#4

Group D Group F Species Av.Abund Av.Abund Av.Diss Diss/SD Contrib% Cum.% Spirobranchus triqueter 0.29 3.46 3.76 7.84 6.65 6.65 Sphaerosyllis bulbosa 1.98 0 2.35 4.44 4.16 10.81 Amphipholis squamata 2.83 1 2.22 2.15 3.92 14.73 Mediomastus fragilis 0.69 2.24 1.85 2.2 3.27 18.01 Moerella donacina 0 1.41 1.68 15.61 2.98 20.98

Table 13 The taxa producing the top 20% of the dissimilarity between G & D.

Groups G & D Average dissimilarity = 53.44 HI02#1

Group G Group D Species Av.Abund Av.Abund Av.Diss Diss/SD Contrib% Cum.% Nematoda 8.72 5.34 3.73 3.8 6.98 6.98 Aonides paucibranchiata 5.48 2.22 3.57 5.96 6.69 13.67 Enchytraeidae 3.61 1.28 2.57 2.55 4.8 18.47 Spirobranchus lamarcki 2.24 0.14 2.31 4.81 4.32 22.79

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4. DISCUSSION

4.1 Study of maerl beds in the South Arran MPA

The South Arran MPA contains expansive areas of coarse sediment biotopes, often containing high percentages of dead maerl and maerl gravel. The distribution of dead and live maerl suggests that live maerl beds may once have extended from the Iron Rock Ledges area to north of Drumadoon Point and from the Pladda area past the eastern shore of Holy Isle to Clauchlands Point (Moore, 2014a) (Figure 12).

Figure 12. Distribution of coarse sediment biotopes with dead and live maerl around the South Arran MPA.

To be able to define ares of maerl beds as an MPA protected feature, it was necessary to engender some distinguishing criteria to split up the habitat continua between maerl beds and maerl gravel areas, which after a review of the relevant data by Colin Moore in 2014, was set at 5% live material (O on the SACFOR scale) for this MPA (Moore, 2014a). To take account of the patchiness and gradation between maerl beds and other coarse gravel habitats, the areas mapped as the ‘maerl bed’ protected feature for the South Arran MPA (SNH, 2014) are surrounded by ‘maerl recovery areas, which perhaps represent areas of historically richer maerl with the greatest likelihood for recovery and suitability for conservation management given the vegetative propagation of maerl (see Moore, 2014a). Contemporary maerl beds have been recorded in depths of 4 - 25 m in the MPA; along the northern coastline of Lamlash Bay, off the north-eastern and southern shores of Holy Isle, along the south coast of Arran at Kildonan and on the south-west coast at Sliddery (Axelsson et al., 2010; Allen, 2013.; Moore, 2014b; Howson, 2014; Munro et al., 2014.). The proportion of live maerl on the majority of these beds is very low (1 - 5 %, Munro et al.,

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2014), with the notable exception of two pockets of rich maerl studied here, where live maerl coverage reaches 30% and 75%. Prior to the 2014 drop down video survey (Morris-Webb, 2015) and the following dive survey described in this report, there were no extant records of dense maerl beds with proportions of live maerl above 20% in the South Arran MPA, or indeed the Clyde Sea area. The two dense beds studied here, off Pladda and in Lamlash Bay, prove that maerl beds in the Clyde Sea area can indeed be made up of large well developed live fragments at high percentages - and thereby create the complex three-dimensional structure which forms the basis of much of the biotope’s diversity and value as a nursery habitat for many marine animals including commercial species of fish and shellfish such as queen scallops (Kamenos et al., 2004). It proves that the Clyde’s environmental regime is indeed suitable for such beds, and lends substantial strength to the argument that much of the maerl gravel around Arran derives from degraded, previously dense, maerl beds. Moore (2014a) noted that the correlation between lower levels of scallop dredging and higher levels of live maerl around the South of Arran, and the presence of dense dead, relatively unbroken maerl and high levels of scallop dredging off the south of Holy Isle, are suggestive, though not proof, of dredging impact. The relationship between scallop dredging and degradation of maerl beds has been confirmed by studies in other locations in the Clyde such as Hall-Spencer and Moore (2000) and Hauton, et al. (2003), and Kamenos et al (2003). The two rich maerl beds mapped in 2014 both lie in areas which are relatively inaccessible to dredgers. The Pladda bed is located in a shallow bay protected by a bank of boulders, and the Lamlash Bay bed is confined to a bend in a steep slope on the north side of the channel between Holy Isle and Arran, which would have been awkward to dredge. Since it is now obvious that the environment in the South Arran MPA is indeed suitable to support full-coverage live beds of well-formed fragments, this is a realistic target for recovery, albeit very long term. Three different maerl habitats were encountered in the South Arran MPA. The bed of Phymatolithon calcareum at Pladda, formed by large medallions of maerl, on the exposed south coast of the island; the maerl gravel areas on the semi-exposed coast of Holy Isle which were also dominated by sparse Phymatolithon calcareum, and the more sheltered maerl bed in Lamlash Bay which was dominated by twigs of Lithothamnion glaciale (see Figure 3 and Figure 7). Since maerl species are difficult to identify to species level without specialised laboratory methods it had not previously been known that the Pladda and Lamlash Bay beds were made up of different species. This finding further emphasises the complexity of the South Arran ecosystem, which contains niches for a variety of diverse biotopes. Of concern, particularly around Holy Isle, are the large amounts of the non-native algal species Dasysiphonia japonica on both maerl beds and ‘Trailliella intricata’ (the tetrasporophyte phase of Bonnemaisonia hamifera) on the live maerl in Lamlash Bay. 4.2 Comparison of the ‘maerl bed’ and ‘maerl gravel’ biotopes

This survey studied two sites on mapped ‘maerl beds’ (PL32 and LBN) and three in areas of coarse sediment north and south of Holy Isle (HIN04 and HIN02 north, and HI22 south). Comparison of the biological diversity of associated communities shows clearly that there is a difference between intact ‘maerl beds’ and maerl gravel areas. Values of all univariate community indices were higher at the two maerl bed sites than at the three coarse sediment sites (e.g. mean Shannon Diversity H’ was 3.09 for LBN and 2.83 for PL32 versus 2.78, 2.43 and 2.70 at the Holy Isle sites). Some of the differences in the composition of the biological communities between the Holy Isle sites and Pladda and Lamlash Bay are caused by the different exposure regimes - sheltered in Lamlash Bay via medium exposure around Holy

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Isle to exposed at Pladda. This is especially true for the epifloral and sessile epibenthic communities. Biological diversity of the infaunal communities, however, seems to be correlated with the presence of live maerl. A comparison of the effects of maerl bed quality on highly mobile epifauna (especially fish) would be highly beneficial to assess the ecosystem value to the MPA, and to describe it in ways meaningful to most people. It is possible that the patches remaining today are too small and isolated to show an effect at that level, but further studies to investigate this are currently underway, such as baited camera studies investigating the use of different habitats around South Arran by juvenile gadoid fish (Elliott et al., 2015). Comparing the two sites north of Holy Isle, which were very close to each other but just either side of the NTZ boundary (HIN04 and HIN02) did not expose meaningful differences which could be ascribed to effects of the NTZ. No activity data exists; however, to confirm to what degree pressures on these spatially very close sites actually differed. The conspicuous dominant taxa were very similar and the two transects appeared to be part of one continuum. It was notable that individual flame shells (Limaria hians) were seen at both of these stations, though a search in the vicinity of the transects did not reveal beds of adults. Given the presence of solitary individual adults and juvenile flame shells, it is possible that flame shell bed patches do exist in the shallows around the coast of Holy Isle. 4.3 Macrobenthic infauna - comparison to other Arran and UK maerl bed records

Previous core data on infaunal diversity of maerl beds around Arran is restricted to a series of single 10.3 cm diameter core samples, collected from three discrete SS.SMp.Mrl.Pcal.R maerl bed survey stations off Clauchlands Point on the northern coastline of Lamlash Bay between 4 - 11 m depth in 2010 (Munro et al., 2014). The 2010 infaunal core samples were assigned to the ‘shallow tide-swept coarse sands with burrowing bivalves’ protected feature biotope (SS.SCS.ICS.MoeVen) on the basis of the key characterising species present within the sediment. Infaunal taxon richness ranged from 35 - 82 taxa (35, 62 and 82) with infaunal abundances between 96 - 724 (96, 372 and 724) ind. / 0.01 m2. These core samples were collected as part of baseline monitoring of the NTZ and were processed using a 0.5 mm mesh sieve, resulting in the representation of a greater number of small invertebrate taxa (especially polychaete worms) than would have been recorded using a 1 mm sieve (Munro et al., 2014). The latter has been used more frequently in recent comparative studies of maerl bed infaunal diversity (e.g. Moore et al., 2011). Because of the very small sample size in 2010 and the methodological differences it is impossible to say whether the differences in taxon richness and abundances reflect actual temporal or spatial trends when compared to the findings or this study (S mean 29 range 21-38, N mean 84 range 35-183). Grab samples collected by SNH in 2013 along the maerl / maerl gravel band south-west off Arran were also assigned to the biotopes SS.SCS.ICS and SS.SCS.ICS.MoeVen (Allen et al., 2014). Infaunal diversity of two single day grabs processed through a 1 mm sieve collected within the maerl beds east of Pladda and west of Sliddery was very high with Shannon Diversity (H’) values of 5.29 and 5.09, and 105 and 77 species (Allen et al., 2014). Again, quantitative comparison with the 2014 core data is complicated by the methodological differences, but creates the impression that the infaunal communities of this survey do not stand out for their diversity and richness. Moore et al. (2011) collated results of 12 similar Scottish maerl bed studies. The infaunal sampling effort in those was slightly smaller than in this study, with only four cores taken per site instead of five. Site-wise infaunal taxon richness described by Moore, of between 56 and 122 species leaves Arran’s maerl beds on the low side with between 50 and 71 taxa, and the

23

mean infaunal abundance values of between 72 and 453 described by Moore compared to Arran’s 56 to 106, also leave Arran slightly on the low side. Further context is provided by studies of beds using similar techniques in the Sound of Arisaig (Moore et al., 2004, Moore et al., 2015), Loch nam Madadh (Moore et al., 2006), Loch Laxford (Moore et al., 2010), and Fetlar to Haroldswick (Hirst et al. 2013). Infaunal taxon richness of cores collected at these sites ranged from 30 to 109 species per sample, which the range of 21 - 38 species in the 2014 Arran cores compares to unfavourably. This may be a reflection of the very small size of the dense maerl bed patches in Arran, which remain in only a few inaccessible corners of a heavily dredged coastline. Diversity indices follow the same trend, with the 2014 Arran beds yielding Shannon Wiener diversities of H’ = 2.43 - 3.09 compared to H’ 2.9 - 5.22 in the above mentioned studies. The numbers of epifaunal and floral taxa recorded on the South Arran maerl beds appear to contradict the depauperate trend above, with Arran’s beds counting between 68 and 89 taxa compared to 33 to 109 taxa on of the other beds from Arisaig (x5), Loch Maddy (x4) and Loch Laxford (x3). On a UK level, compared to a maerl bed in Milford Haven, South Wales, the South Arran 2014 samples rank quite favourably. One hundred and thirty taxa were recorded overall in the 25 cores taken at the five Arran dive sites. Milford Haven, for a similar sampling effort, over five sampling occasions, returned between 86 and 142 taxa in a similar 25 cores, with an overall mean of 107 taxa per sampling occasion. This indicates the importance Scotland plays for maerl beds on a UK level, with many excellent examples found here, but also puts into perspective the ‘poorer’ Arran beds, which are still quite rich when compared to beds further south. Of interest in the macrobenthic cores were the records of small flame shell L. hians specimens at both HIN04 and LBN. These core records supplement the diver observations of individuals of this species at HIN04 and HIN02 (see also Section 4.2). 4.4 Whiting Bay seagrass bed

The diver survey of the seagrass bed in Whiting Bay revealed that the bed, while patchy, compares very favourably with other seagrass beds on the Scottish west coast. Diver MNCR phase 2 style surveys of two beds in Wester Ross using a similar methodology yielded average shoot densities of 46/m2 with 49 epibenthic taxa, and 113/m2 with 56 epibenthic taxa (Moore et al. 2011), compared to the 89 shoots m-2 and 60 epibenthic taxa recorded here. The abundant echiuran(?) seen in the sediments within the Echinocardium caudatum beds perhaps warrants further investigation.

24

5. REFERENCES

Allen, J.H. 2013. Infaunal analysis of grab samples collected from the Clyde Sea, in March 2012. Scottish Natural Heritage Commissioned Report No. 539.

Allen, C., Axelsson, M. & Dewey, S. 2014. Marine biological survey to establish the distribution of Priority Marine Features within the Clyde Sea area. Scottish Natural Heritage Commissioned Report No. 437.

Axelsson, M., Dewey, S., Doran, J. & Plastow, L. 2010. Mapping of marine habitats and species of Lamlash Bay, Arran. Scottish Natural Heritage Commissioned Report No. 400. Elliott, S.A.M, Sabatino, A.D., Heath,M.R., Turrell, W.R. & Bailey, D.M. 2015. Predictive substratum modelling for juvenile gadoid distribution and abundance. Conference Paper, ICES CM 2015/N:09. Hall-Spencer, J.M. and Moore, P.G. 2000. Scallop dredging has profound, long-term impacts on maerl habitats. ICES Journal of Marine Science, 57, 1407-1415. Hauton, C., Hall-Spencer, J.M. & Moore, P.G. 2003. An experimental study of the ecological impacts of hydraulic bivalve dredging on maerl. ICES Journal of Marine Science, 60, 381-392. Hirst, N.E., Kamphausen, L.M., Cook, R.L., Porter, J.S. & Sanderson, W.G. 2013. The distribution and status of proposed protected features within the Fetlar and Haroldswick MPA proposal. Scottish Natural Heritage Commissioned Report 599. Holme, N.A. & McIntyre, A.D. 1984. Eds. Methods for the Study of Marine Benthos. Second Edition, IBP Handbook 16. 399 pp. Oxford-London-Boston: Blackwell Scientific Publications 1984. ISBN 0-632-00894-6. Howson, C. & Steel, L. 2014. Validation of seabed habitat MPA search feature records relating to the South Arran Nature Conservation MPA. Scottish Natural Heritage Commissioned Report No. 620. Howson, C.M. & Picton, B.E. eds. 1997. The species directory of the marine fauna and flora of the British Isles and surrounding seas. Ulster Museum and The Marine Conservation Society, Belfast and Ross-on-Wye. Belfast: Ulster Museum. Ulster Museum Publication No. 276. JNCC, 1990. SACFOR abundance scale used for both littoral and sublittoral taxa from 1990 onwards. http://jncc.defra.gov.uk/page-2684 Kamenos, N.A., Moore, P.G., and Hall-Spencer, J.M. 2004. Nursery-area function of maerl grounds for juvenile queen scallops Aequipecten opercularis and other invertebrates. Marine Ecological Progress Series, 274, 183-189. Kamenos, N.A., Moore P.G. & Hall-Spencer, J.M. 2003. Substratum heterogeneity of dredged vs. un-dredged maerl grounds. Journal of Marine Biological Association, 83, 411-413. Marine Scotland. 2014. Consultation on the management of inshore Special Areas of Conservation and Marine Protected Areas - Overview. http://www.gov.scot/Resource/0046/00462816.pdf

25

Moore, C.G. 2014a. The distribution of maerl and other coarse sediment proposed protected features within the South Arran pMPA - a data review to inform management options. Scottish Natural Heritage Commissioned Report No. 749. https://www.nature.scot/snh-commissioned-report-749-distribution-maerl-and-other-coarse-sediment-proposed-protected-features Moore, C.G. 2014b. Upper Loch Fyne and Loch Goil pMPA and Wester Ross pMPA - the identification of conservation management areas to support protected feature recovery. Scottish Natural Heritage Commissioned Report No. 764. Moore, C.G., Harries, D.B., Cook, R.L., Saunders, G.R., Atkinson, R.J.A. & Sanderson, W.G. 2015. 2014 site condition monitoring survey of marine sedimentary habitats in the Sound of Arisaig SAC. Scottish Natural Heritage Commissioned Report No. 807. https://www.nature.scot/snh-commissioned-report-807-2014-site-condition-monitoring-survey-marine-sedimentary-habitats-sound Moore, C.G., Harries, D.B., Trigg, C., Porter, J.S. & Lyndon, A. R. 2011. The distribution of Priority Marine Features and MPA search features within the Ullapool Approaches: a broadscale validation survey. Scottish Natural Heritage Commissioned Report No. 422. https://www.nature.scot/snh-commissioned-report-422-distribution-priority-marine-features-and-mpa-search-features-within Moore, C.G., Harries, D.B., Porter, J.S. & Lyndon, A.R. 2010. The establishment of site condition monitoring of marine features of Loch Laxford Special Area of Conservation. Scottish Natural Heritage Commissioned Report No. 378. https://www.nature.scot/snh-commissioned-report-378-establishment-site-condition-monitoring-marine-features-loch-laxford Moore, C.G, Saunders, G., Mair J.M. & Lyndon A.R. 2006. The inauguration of site condition monitoring of marine features of Loch Maddy Special Area of Conservation. Scottish Natural Heritage Research Commissioned Report No. 152. Moore, C.G., Lyndon, A.R. & Mair, J.M. 2004. The establishment of site condition monitoring of marine sedimentary habitats in the Sound of Arisaig cSAC. Scottish Natural Heritage Commissioned Report No.071 (ROAME No. F02AA409). Morris-Webb, E.S. 2015. Biological analyses of underwater video footage from Arran, Loch Linnhe, Loch Shell and Loch Seaforth. Scottish Natural Heritage Commissioned Report No. 818. https://www.nature.scot/snh-commissioned-report-818-biological-analyses-underwater-video-footage-arran-loch-linnhe-loch Munro, C.D., Baldock, L., Brown, K. and Lindsley-Leake, S. (2014). Lamlash Bay, Arran, 2010 survey report. Scottish Natural Heritage Commissioned Report No. 619. Scottish Statutory Instruments, 2015. The South Arran Marine Conservation Order 2015. http://www.legislation.gov.uk/ssi/2015/437/made SNH. 2014. South Arran Nature Conservation MPA. Scottish MPA Project - Assessment against the MPA Selection Guidelines. https://www.nature.scot/professional-advice/safeguarding-protected-areas-and-species/protected-areas/national-designations/marine-protected-areas/nature-conservation-13

26

ANNEX 1: RAW DATA

Table A. Site characteristic data for all dive sites

Site name Pladda Holy Isle

South Holy Isle

North Lamlash

Bay North Holy Isle

North Whiting

Bay Site code PL34 HI22 HIN04 LBN HIN02 WBZ

Survey date 21/09/2014 22/09/2014 23/09/2014 24/09/2014 25/09/2014 26/09/2014 Latitude 55.43635 55.51077 55.53867 55.54600 55.53563 55.49758 Longitude -5.12493 -5.06212 -5.07758 -5.08857 -5.07232 -5.08833 Transect bearing 240 220 140 100 320 n/a Personnel Phase 2 survey FB/JM FB/JM FB/JM FB/JM FB/JM FB/JM Coring TM TM TM TM TM n/a Maerl/seagrass quadrats

LK/JAT LK/JAT LK/JAT LK/JAT LK/JAT LK/JAT

Photography FB/JAT/LK FB/JAT/LK FB/JAT/LK FB/JAT/LK FB/JAT/LK FB/JAT/LK Video LK TM TM TM TM TM

Substrata % Bedrock / boulders 0 0 0 0 0 0 Cobbles 0.5 0.1 0 0 0 0 Whole shell 30 18.78 23.85 15 20 5 Pebbles 5.5 0.1 0.1 0 1 2 Live maerl 20 0.01 0.05 55 1 0 Maerl gravel 30 80 70 25 74 0 Gravel 10 0 0 0 0 1 Sand 4 1 3 0 2 92 Mud 0 0.01 3 5 2 0 Total (calc) 100 100 100 100 100 100

Depth BCD (m) 7.5 7.8 6.2 9.8 6 4.4

Sediment features (1 - 5) Surface relief (even-uneven)

2 2 3 2 2 1

Firmness (firm-soft) 3 3 3 4 3 3 Stability (stable-mobile) 3 3 3 2 3 2 Sorting (well-poor) 4 3 4 3 3 4

Sediment features (presence) Mounds / casts N N N N N N Burrows / holes N Y N Y N Y Tubes Y Y Y Y Y Y Algal mat N N Y Y Y N Waves / dunes (>10 cm high)

N N N N N N

Ripples (<10 cm high) N N N N N N Subsurface black layer N N N ? N N Subsurface coarse layer

N N N N N N

Subsurface clay / mud N N Y Y Y N Surface silt / flocculent N N N N N N

27

Table B. Quadrat records of Site Maerl Abundance

Site PL34 21/09/2014 HI22 22/09/2014 HIN04 23/09/2014 LBN 24/09/2014 HIN02 25/09/2014

Quadrat % live maerl

% total maerl / maerl gravel diver

% live maerl


% live maerl


% live maerl


% live maerl


L1 30 95 LK 1 98 LK 1 100 LK 85 95 LK 0.5 100 LK

L2 25 95 LK 2 100 LK 3 100 LK 80 95 LK 1 100 LK

L3 15 90 LK 1 99 LK 3 100 LK 80 95 LK 0.5 95 LK

L4 40 95 LK 2 95 LK 1 95 LK 90 100 LK 0.5 100 LK

L5 20 95 LK 2 95 LK 2 100 LK 70 90 LK 0 95 LK

L6 30 95 LK 2 100 LK 1 100 LK 80 85 LK 0.5 100 LK

L7 20 90 LK 1 98 LK 1 95 LK 80 95 LK 0 100 LK

L8 15 90 LK 1 100 LK 2 100 LK 80 95 LK 0.5 100 LK

L9 45 90 LK 5 100 LK 3 100 LK 70 80 LK 1 100 LK

L10 30 95 LK 1 100 LK 2 100 LK 50 90 LK 1 100 LK

R1 60 90 JAT 5 100 JAT 2 100 JAT 50 95 JAT 1 100 JAT










Mean 28.75 92.25 2.3 97.15 2.4 98.5 75.5 90.5 0.925 99.25

28

Table C. Abundance (SACFOR) data for the fauna and flora of the maerl and Zostera biotopes

Scientific name accepted

Sp Code AphiaID accepted

Authority accepted

Notes Spec.

checked Photos PL34 HI22 HIN04 LBN HIN02 WBZ

No. of recorded taxa 193 68 83 86 89 85 60 Porifera (white 'Pennar' sponge)

C000001 558 Grant, 1836 R R

Suberites C002180 132072 Nardo, 1833 A few small colonies R R R Halichondria bowerbanki C004810 165801 Burton, 1930 Y (not

spicules) x1

Amphilectus fucorum C005960 150225 (Esper, 1794) A couple of small colonies

R

Haliclona C008540 131834 Grant, 1836 Y R R Hydractinia echinata D003350 117644 (Fleming, 1828) Y R R Sertularella gayi D006670 117902 (Lamouroux,

1821) A couple of small colonies on shells

R

Clytia hemisphaerica D007030 117368 (Linnaeus, 1767)

On Polysiphonia elongata

Y P

Obelia sp. D007280 117034 Péron & Lesueur, 1810

Y R

Obelia dichotoma D007300 117386 (Linnaeus, 1758)

R R

Obelia geniculata D007310 117388 (Linnaeus, 1758)

On mature Saccharina latissima fronds

R R R R

Cerianthus lloydii D010750 283798 Gosse, 1859 Y F A F R O R Sagartiogeton laceratus D012470 100999 (Dalyell, 1848) O Sagartiogeton undatus D012480 101002 (Müller, 1778) Y Y O R Lineus longissimus G000780 122528 (Gunnerus,

1770) R

Echiurus echiurus O000040 110377 (Pallas, 1766) Long mottled orangey pink proboscis (3 mm wide, blunt end) extending for up to 10" from burrow, creating radial marks around burrow

C

29

Polynoidae P000420 939 Kinberg, 1856 P Alentia gelatinosa P000600 130722 (M. Sars, 1835) Y P Lepidonotus squamatus P001330 130801 (Linnaeus,

1758) Y P

Chaetopterus P013720 129229 Cuvier, 1830 Y F F O C C O Terebellidae P020000 982 Johnston, 1846 x1 x1 Myxicola infundibulum P022270 130932 (Montagu, 1808) R R R Hydroides norvegicus P022880 131009 Gunnerus, 1768 Y Y R C O R Spirobranchus P023020 129582 Blainville, 1818 Y F O F C F O Spirobranchus lamarcki P023030 560033 (Quatrefages,

1866) Y P

Spirobranchus triqueter P023040 555935 (Linnaeus, 1758)

Y P P P P

Spirorbinae P023550 989 Chamberlin, 1919

Y O O O C O

Balanus balanus R001090 106213 (Linnaeus, 1758)

Y P

Balanus crenatus R001100 106215 Bruguière, 1789 Y Y P R Ericthonius punctatus S009440 102408 (Bate, 1857) Y P Pandalus montagui S023220 107651 Leach, 1814 [in

Leach, 1813-1814]

Y O x1

Anapagurus hyndmanni S024470 107217 (Bell, 1846) R O x1 Pagurus bernhardus S024650 107232 (Linnaeus,

1758) Y R O O O C

Pagurus cuanensis S024680 107235 Bell, 1846 x1 Galathea sp. (juv) S024840 106834 Fabricius, 1793 P Galathea intermedia S024860 107150 Liljeborg, 1851 Y Y C Munida rugosa S024950 107160 (Fabricius,

1775) x1

Pisidia longicornis S025020 107188 (Linnaeus, 1767)

P P

Hyas araneus S025590 107322 (Linnaeus, 1758)

x1 x1

Inachus dorsettensis S025760 107327 (Pennant, 1777) Y Y O C x1

30

Macropodia rostrata S025850 107345 (Linnaeus, 1761)

Y O O x1 C

Cancer pagurus S026460 107276 Linnaeus, 1758 Y O O F O F Liocarcinus corrugatus S026680 107386 (Pennant, 1777) Y x1? Liocarcinus depurator S026690 107387 (Linnaeus,

1758) x1 O C

Necora puber S026720 107398 (Linnaeus, 1767)

Y F F F C F F

Carcinus maenas S026900 107381 (Linnaeus, 1758)

O

Polyplacophora W000500 55 Gray, 1821 On dead shells P P Polyplacophora (patterned)

W000500 55 Gray, 1821 On dead shells P

Leptochiton cancellatus W000560 140201 (Sowerby, 1840) On dead shells Y P P Lepidochitona cinerea W000740 140143 (Linnaeus,

1767) On dead shells Y P

Testudinalia testudinalis W001250 234208 (O. F. Müller, 1776)

On dead shells Y F P P P

Tectura virginea W001260 153552 (O. F. Müller, 1776)

On dead shells Y F C

Gibbula magus W001890 141790 (Linnaeus, 1758)

Y Y O F R O

Lacuna vincta W002440 140170 (Montagu, 1803) Y P P Rissoa lilacina W002810 141358 Récluz, 1843 Y P P P P C Rissoa parva W002850 141365 (da Costa,

1778) Y P C

Trivia arctica W007370 141741 (Pulteney, 1799) Y x1 Buccinum undatum W008440 138878 Linnaeus, 1758 x1 Aplysia punctata W011020 138758 (Cuvier, 1803) Y x1 P P Onchidoris pusilla W013370 140643 (Alder &

Hancock, 1845) Y P P

Limacia clavigera W013580 140830 (O. F. Müller, 1776)

Y R P

Limaria hians W017390 140235 (Gmelin, 1791) Y R R Chlamys varia W018000 236719 (Linnaeus,

1758) Y R

31

Aequipecten opercularis W018050 140687 (Linnaeus, 1758)

Mostly juveniles Y R O O

Pecten maximus W018090 140712 (Linnaeus, 1758)

Y O x1 F

Pecten maximus (juv) W018090 140712 (Linnaeus, 1758)

R F

Pododesmus patelliformis

W018200 153027 (Linnaeus, 1761)

Y O

Ensis sp.(siphons) W020220 138333 Schumacher, 1817

O O

Ensis arcuatus W020230 160539 Schumacher, 1817

C

Dosinia exoleta W021660 141911 (Linnaeus, 1758)

Y F P

Polititapes rhomboides W021810 745846 (Pennant, 1777) Dead shells common Y R F O Chamelea gallina W021890 141907 (Linnaeus,

1758) x1

Crisiidae Y000030 110806 Johnston, 1838 R Crisia denticulata Y000270 111695 (Lamarck, 1816) Y R Crisia eburnea Y000280 111696 (Linnaeus,

1758) Y R

Tubulipora plumosa Y000530 111765 Thompson in Harmer, 1898

Frequent on some mature Saccharina latissima fronds

Y P R

Fenestrulina malusii Y005230 111418 (Audouin, 1826) Common on dead Dosinia shells

Y R R

Celleporella hyalina Y005710 111397 (Linnaeus, 1767)


Y Y O O O O O O

Aetea truncata Y006450 111067 (Landsborough, 1852)

On Phyllophora crispa Y P R

Membranipora membranacea

Y006640 111411 (Linnaeus, 1767)


O O O O O

Electra pilosa Y006780 111355 (Linnaeus, 1767)


Y O O O O O

Cradoscrupocellaria reptans

Y008380 738997 (Linnaeus, 1758)

Common on Desmarestia aculeata plants

Y Y R R R R R R

32

Scrupocellaria scrupea Y008400 111249 Busk, 1852 Y R Scrupocellaria scruposa Y008410 111250 (Linnaeus,

1758) On dead bivalve shell Y R R

Antedon bifida ZB00110 124201 (Pennant, 1777) R Astropecten irregularis ZB00410 123867 (Pennant, 1777) Y x1 x1 O x1 Luidia ciliaris ZB00670 123920 (Philippi, 1837) Y x1 F O x1 Porania pulvillus ZB01010 125166 (O.F. Müller,

1776) Y F O O

Crossaster papposus ZB01490 124154 (Linnaeus, 1767)

Y x1 O x1

Henricia sp. ZB01640 123276 Gray, 1840 Y O Asterias rubens (juv) ZB01900 123776 Linnaeus, 1758 Y O O O O O Asterias rubens ZB01900 123776 Linnaeus, 1758 Incl. some enormous

flabby individuals Y O C O x1

Marthasterias glacialis ZB02000 123803 (Linnaeus, 1758)

Y C O F O F

Ophiothrix fragilis ZB02350 125131 (Abildgaard, in O.F. Müller, 1789)

Y P R P

Amphipholis squamata ZB03000 125064 (Delle Chiaje, 1828)

Y P

Ophiura albida ZB03130 124913 Forbes, 1839 Y P Ophiura ophiura ZB03150 124929 (Linnaeus,

1758) x1 F

Echinoidea (juv) ZB03380 123082 Leske, 1778 In dead bivalve shells Y P Echinus esculentus ZB03620 124287 Linnaeus, 1758 x1 Echinocardium cordatum

ZB04070 124392 (Pennant, 1777) C

Clavelina lepadiformis ZD00060 103552 (Müller, 1776) R R Aplidium punctum ZD00640 103662 (Giard, 1873) Y R Diplosoma listerianum ZD00970 103579 (Milne-Edwards,

1841) Particularly on mature Saccharina latissima fronds

Y R O R R R

Ciona intestinalis ZD01170 103732 (Linnaeus, 1767)

Mostly hidden inside valves of dead bivalve shells

R R R

33

Corella parallelogramma ZD01350 103743 (Müller, 1776) R O O Ascidiella aspersa ZD01410 103718 (Müller, 1776) Y Y O F O O C O Ascidiella scabra ZD01430 103719 (Müller, 1776) Y R Botryllus schlosseri ZD02090 103862 (Pallas, 1766) R Scyliorhinus canicula ZF00400 105814 (Linnaeus,

1758) x1

Diplecogaster bimaculata

ZG01240 236458 (Bonnaterre, 1788)

Y x1

Agonus cataphractus ZG04480 127190 (Linnaeus, 1758)

x1

Ctenolabrus rupestris ZG06050 126964 (Linnaeus, 1758)

P

Pholis gunnellus ZG06800 126996 (Linnaeus, 1758)

O O x1 F

Callionymus reticulatus ZG07020 126795 Valenciennes, 1837

Y O O O O

Gobiusculus flavescens ZG07280 126898 (Fabricius, 1779)

x1

Pomatoschistus ZG07410 125999 Gill, 1863 Y C F F F F C Limanda limanda (juv) ZG08910 127139 (Linnaeus,

1758) x1

Pleuronectes platessa ZG09030 127143 Linnaeus, 1758 R Rhodophyta (enc) ZM00010 852 Wettstein, 1901 On pebbles R R R R R Acrochaetiaceae ZM00960 143661 Fritsch ex W.R.

Taylor, 1957 Y R R

Scinaia interrupta ZM01820 146401 (A.P.de Candolle) M.J.Wynne, 1989

Y Y F O O R O

Asparagopsis armata (Falkenbergia rufolanosa)

ZM02020 144438 Harvey, 1855 R R

Bonnemaisonia asparagoides

ZM02080 144440 (Woodward) C.Agardh, 1822

R

Bonnemaisonia hamifera (Trailliella intricata)

ZM02110 144442 Hariot, 1891 Y Y R S R

34

Palmaria palmata ZM02420 145771 (Linnaeus) Weber & Mohr, 1805

Y R R

Dilsea carnosa ZM02560 145222 (Schmidel) Kuntze, 1898

Y R

Dudresnaya verticillata ZM02610 145226 (Withering) Le Jolis, 1863

R

Callophyllis laciniata ZM03230 145262 (Hudson) Kützing, 1843

Y R

Peyssonnelia ZM03640 144051 Decaisne, 1841 R R R R Corallinaceae (enc) ZM03840 143691 Lamouroux,

1812 Y O O O O O R

Corallina officinalis ZM04040 145108 Linnaeus, 1758 Y R O Lithothamnion glaciale ZM04610 145170 Kjellman, 1883 R S Phymatolithon calcareum

ZM04910 145199 (Pallas) W.H.Adey & D.L.McKibbin, 1970

Y C R R O

Pneophyllum fragile ZM05050 145206 Kützing, 1843 Y R Coccotylus truncatus ZM05830 145654 (Pallas)

M.J.Wynne & J.N.Heine, 1992

Y R

Phyllophora crispa ZM05840 145660 (Hudson) P.S.Dixon, 1964

Y R R

Erythrodermis traillii ZM05880 145655 (Holmes ex Batters) Guiry & Garbary, 1990

Y R

Chondrus crispus ZM06110 145625 Stackhouse, 1797

Y R

Polyides rotunda ZM06250 145668 (Hudson) Gaillon, 1828

R

Rhodophyllis divaricata ZM06930 145617 (Stackhouse) Papenfuss, 1950

Y R R R R R

Cruoria cruoriiformis ZM07010 145611 (P.L.Crouan & H.M.Crouan) Denizot, 1968

Y R? R? R? R? R?

35

Chylocladia verticillata ZM07400 145808 (Lightfoot) Bliding, 1928

R R

Lomentaria clavellosa ZM07520 145825 (Lightfoot ex Turner) Gaillon, 1828

Y R R R R R R

Lomentaria orcadensis ZM07530 145830 (Harvey) F.S.Collins, 1937

R

Aglaothamnion bipinnatum

ZM07850 144489 (P.L.Crouan & H.M.Crouan) Feldmann & G.Feldmann, 1948

Y P

Callithamnion corymbosum

ZM07890 144526 (Smith) Lyngbye, 1819

R

Ceramium secundatum ZM08239 144562 Lyngbye, 1819 Y R R R Compsothamnion thuyoides

ZM08340 144573 (Smith) Nägeli, 1862

Y R

Halurus flosculosus ZM08460 144595 (J.Ellis) Maggs & Hommersand, 1993

R

Pterothamnion plumula ZM08880 144683 (J.Ellis) Nägeli, 1855

Y R R R

Ptilota gunneri ZM08930 144686 P.C.Silva, Maggs & L.M.Irvine, 1993

Y R

Ptilothamnion sphaericum

ZM08985 144690 (P.L.Crouan & H.M.Crouan ex J.Agardh) Maggs & Hommersand, 1993

Y R

Seirospora interrupta ZM09080 144697 (Smith) F.Schmitz, 1893

Y R

Spermothamnion repens ZM09170 144702 (Dillwyn) Rosenvinge, 1924

Y R R R R

36

Spermothamnion strictum

ZM09180 144704 (C.Agardh) Ardissone, 1883

Y R R R

Sphondylothamnion multifidum

ZM09230 144705 (Hudson) Nägeli, 1862

Y R R

Apoglossum ruscifolium ZM09400 144737 (Turner) J.Agardh, 1898

R R R

Cryptopleura ramosa ZM09500 144743 (Hudson) L.Newton, 1931

Y R R R

Delesseria sanguinea ZM09550 144744 (Hudson) J.V.Lamouroux, 1813

Y R R R R

Hypoglossum hypoglossoides

ZM09850 144756 (Stackhouse) F.S.Collins & Hervey, 1917

Y R R

Membranoptera alata ZM09900 144758 (Hudson) Stackhouse, 1809

R

Haraldiophyllum bonnemaisonii

ZM09950 144755 (Kylin) A.D.Zinova, 1981

R

Nitophyllum punctatum ZM10020 144770 (Stackhouse) Greville, 1830

Y R R

Heterosiphonia japonica ZM10380 232226 Yendo, 1920 Y Y F F S O C F Heterosiphonia plumosa ZM10390 144732 (J.Ellis) Batters,

1902 Y R R R R

Chondria dasyphylla ZM10560 144799 (Woodward) C.Agardh, 1817

Y R

Odonthalia dentata ZM10970 144839 (Linnaeus) Lyngbye, 1819

Y R

Polysiphonia elongata ZM11050 144628 (Hudson) Sprengel, 1827

Y R R P R R R

Polysiphonia fucoides ZM11170 144639 (Hudson) Greville, 1824

Y R R R

Polysiphonia stricta ZM11300 144672 (Dillwyn) Greville, 1824

Y R R R R

Pterosiphonia parasitica ZM11370 144851 (Hudson) Falkenberg, 1901

Y R R R R R

37

Rhodomela confervoides

ZM11450 144854 (Hudson) P.C.Silva, 1952

Y R R

Bacillariophyceae ZN 148899 Y R Cutleria multifida ZR03890 145297 (Turner)

Greville, 1830 Y R

Cutleria multifida (Aglaozonia)

ZR03890 145297 (Turner) Greville, 1830

Common on dead bivalve shells

Y R R R O O R

Sphacelaria ZR04120 144272 Lyngbye, 1818 Y R R R R Dictyota dichotoma ZR04570 145367 (Hudson)

J.V.Lamouroux, 1809

Y R R R R R

Desmarestia aculeata ZR04970 145307 (Linnaeus) J.V.Lamouroux, 1813

Y O F R R R R

Desmarestia ligulata ZR04990 145309 (Stackhouse) J.V.Lamouroux, 1813

R

Striaria attenuata ZR05370 548021 (Greville) Greville, 1828

Y R?

Colpomenia peregrina ZR06050 145856 Sauvageau, 1927

Found on 1 dead shell Y R

Chorda filum ZR06250 145722 (Linnaeus) Stackhouse, 1797

Y R R R

Saccharina latissima ZR06360 234483 (Linnaeus) C.E.Lane, C.Mayes, Druehl & G.W.Saunders, 2006

Y F C C F C F

Saccorhiza polyschides ZR06460 145735 (Lightfoot) Batters, 1902

Y O

Halidrys siliquosa ZR07160 145540 (Linnaeus) Lyngbye, 1819

R

Chlorophyta (fluff) ZS00001 801 Pascher, 1914 R Ulva clathrata ZS02140 156078 (Roth)

C.Agardh, 1811 Y R

38

Ulva (flat) ZS02400 144296 Linnaeus, 1753 O R R R R R Ulva (tubular) ZS02400 144296 Linnaeus, 1753 R Chaetomorpha aerea ZS03310 145020 (Dillwyn)

Kützing, 1849 Y R

Cladophora (fine) ZS03380 143996 Kützing, 1843 Y R Derbesia ZS03960 143814 Solier, 1846 Y R R R Zostera marina ZZ 145795 Linnaeus, 1753 A

39

Table D. Microscopic analysis of 10 cm2 maerl samples

Site PL34.A

PL34.B

PL34.C

PL34.D

PL34.E

HI22.A

HI22.B

HI22.C

HI22.D

HI22.E

HI04.A

HI04.B

HI04.C

HI04.D

HI04.E

LBN..A

LBN..B

LBN..C

LBN..D

LBN..E

HI02.A

HI02.B

HI02.C

HI02.D

HI02.E

Algae


P

Acrosorium venulosum

P

P P

P

P

P

Aglaothamnion byssoides

P P

Aglaozonia sp. P P P

Apoglossum ruscifolium

P

P

P

P

Audouinella sp. P

Ceramium secundatum

P

P

Cladophora sp. P

Corallina officinalis P P P P

Coralline crusts P P P P P P P P P P P P P P P P P P P P P P P P P

Cryptopleura ramosa

P P

P P

P

P

P

Cystoclonium purpureum

P P

Dictyota dichotoma P P P P P P P P P P

Ectocarpaceae P

Falkenbergia P P P P

Fine green threads P P P P P P P P P P P

Gelidiella calcicola P P P P P P P P

Griffithsia corallinoides

P

Halarachnion ligulatum

P P

Halurus flosculosus P P

Heterosiphonia japonica

P

P P P P P P P P P P P P P P

P P P P P

P P

40

Site PL34.A

PL34.B

PL34.C

PL34.D

PL34.E

HI22.A

HI22.B

HI22.C

HI22.D

HI22.E

HI04.A

HI04.B

HI04.C

HI04.D

HI04.E

LBN..A

LBN..B

LBN..C

LBN..D

LBN..E

HI02.A

HI02.B

HI02.C

HI02.D

HI02.E

Heterosiphonia plumosa

P

Hypoglossum hypoglossoides

P P

P

Lomentaria clavellosa

P

P P

P

P

P

Phyllophora sp. juv P P P P

Plocamium cartilagineum

P

P P

P

P

Polysiphonia elongata

P

P P

P

P

P

P

Polysiphonia fucoides

P

P P

P

P

P P

P

P P P P P

Polysiphonia harveyi

P

P P

P P P

P P

P

Polysiphonia stricta P P P P P

Pterosiphonia parasitica

P

P P P P P

P

P P P P P P

P

P

Pterothamnion plumula

P

P

P

P

P

P

Ptilothamnion sphaericum

P

P

Ptilota gunneri P

Red crusts P P P P P P P P P P P P P P P P P P P P P P P P P

Rhodomela confervoides

P

P P P

P P

P P

P

P

Rhodophyllis divaricata

P

P P P

P P

P P

P P P P

Scinaia interrupta P P P P

Spermothamnion repens

P P

P P P P P P P P

P P P P P

P

Spermothamnion strictum

P P P

P P P P P P P P

P

P P

P

P P P

Sphacelaria plumosa

P

P

P

P P

P

Sphacelaria sp. P P P P P P P P


P

P

41

Site PL34.A

PL34.B

PL34.C

PL34.D

PL34.E

HI22.A

HI22.B

HI22.C

HI22.D

HI22.E

HI04.A

HI04.B

HI04.C

HI04.D

HI04.E

LBN..A

LBN..B

LBN..C

LBN..D

LBN..E

HI02.A

HI02.B

HI02.C

HI02.D

HI02.E

Trailliella intricata P P P P P P

Ulva lactuca P P

Ulva sp. (flat) P P P P

Ulva sp. (tubular) P

Maerl dead P P P P P P P P P P P P P P P P P P P P P P P P P

Maerl live P P P P P P P P P P P P P P P P P P P

Animals

Ascidiella aspersa P

Ciona intestinalis P

Crisidia cornuta P

Echinus esculentus juv (present in most samples)

P

P P

Fenestrulina malusii

P

Gibbula magus P

Lepidochitona cancellata (present in most samples)

P P P

P

P

Plagioecia patina P

Psammechinus miliaris juv.

P P

Scrupocellaria scruposa

P

Tubulipora liliacea P

Taxa 12 12 16 18 13 11 17 14 10 15 25 20 12 19 17 14 13 15 12 14 18 12 17 14 12

Mean 14.2 13.4 18.6 13.6 14.6

42

Table E. Zostera marina shoot density

Mean density m-2 = 89.30 shoots (quadrat = 0.25 m2)

Quadrat 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 no. of shoots

21 23 26 24 24 18 24 23 24 27 18 12 20 20 24 22 23 24 15 21 22 17 20 28

Quadrat 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 Mean no. of shoots

22 18 15 21 24 34 20 11 20 15 20 18 32 28 35 28 36 34 16 25 20 15 22.33

43

Table F. Maerl InFAUNA core site Particle Size Distribution analyses

Arran maerl core PSD Size PL34 HI22 HIN04 LBN HIN02

Medium pebble (gravel) > 8 mm 9.94 0.00 2.31 0.99 0.00 Small pebble (gravel) 4-8 mm 14.43 4.89 7.47 7.71 0.43 Granule 2-4 mm 38.29 33.61 45.88 20.44 11.91 Sand - very coarse 1-2000 µm 33.89 39.21 39.41 37.86 68.25

Sand - coarse 500-1000

µm 2.01 9.28 1.32 9.49 11.15

Sand - medium 250-500

µm 0.06 4.36 0.20 2.74 2.14

Sand - fine 125-250

µm 0.01 1.58 0.06 0.62 0.59 Sand - very fine 63-125 µm 0.03 1.15 0.12 3.41 0.71 Silt & Clay < 63 µm 1.31 5.92 3.23 16.74 4.82 % Organic Matter (by LOI) 1.75 2.67 2.78 3.12 2.57

44

Table G. Macrobenthic infaunal taxa from Infaunal cores

AphiaID Accepted scientific name

Authority accepted Notes PL34

#1 PL34

#2 PL34

#3 PL34

#4 PL34

#5

Cnidaria

283798 Cerianthus lloydii Gosse, 1859

100665 Edwardsiidae Andres, 1881

Platyhelminthes

793 Platyhelminthes

Nemertea

152391 Nemertea 1 1

Nematoda

799 Nematoda 13 24 14 26 8

Sipuncula

2032 Golfingiidae Stephen & Edmonds, 1972

Juvs 1

410724 Golfingia vulgaris (de Blainville, 1827)

136060 Nephasoma minutum (Keferstein, 1862) ? 1 3 2

Annelida

130707 Pisione remota (Southern, 1914) 1 5 10 7 2

939 Polynoidae Kinberg, 1856 scale-less/juvs 4 5 3 2 3

130770 Harmothoe impar (Johnston, 1839)

130818 Malmgreniella mcintoshi

(Tebble & Chambers, 1982)

1

147008 Malmgrenia andreapolis

McIntosh, 1874

1

130816 Malmgreniella lunulata (Delle Chiaje, 1830) 1 3 2

130599 Pholoe baltica Örsted, 1843

130601 Pholoe inornata Johnston, 1839

931 Phyllodocidae Örsted, 1843 damaged 1

130616 Eteone longa (Fabricius, 1780)

130683 Pseudomystides limbata

(Saint-Joseph, 1888)

130631 Eulalia mustela Pleijel, 1987

129446 Eumida Malmgren, 1865

130641 Eumida bahusiensis Bergstrom, 1914 ?

130644 Eumida sanguinea (Örsted, 1843) 1 2

129296 Glycera Savigny, 1818 juvs

130123 Glycera lapidum Quatrefages, 1866 1 3 1 1

131083 Ephesiella peripatus (Claparède, 1863)

130197 Podarkeopsis helgolandicus

(Hilbig & Dittmer, 1979)

152249 Psamathe fusca Johnston, 1836

130185 Nereimyra punctata (Müller, 1788)

131288 Eurysyllis tuberculata Ehlers, 1864 1

157583 Syllis cornuta Rathke, 1843 1 2 8 5

131458 Syllis variegata Grube, 1860 1

335151 Trypanosyllis coeliaca Claparède, 1868 6 3 1

45



#1 PL34

#2 PL34

#3 PL34

#4 PL34

#5

131379 Sphaerosyllis bulbosa Southern, 1914 2 2 1 2

130417 Platynereis dumerilii (Audouin & Milne Edwards, 1834)

129838 Pareurythoe borealis (M. Sars, 1862) 1

130467 Nothria conchylega (Sars, 1835)

742232 Lysidice unicornis (Grube, 1840)

607402 Hilbigneris gracilis (Ehlers, 1868) spp complex

130041 Protodorvillea kefersteini

(McIntosh, 1869)

8 1 3 4

130044 Schistomeringos neglecta

(Fauvel, 1923)

1

130585 Paradoneis lyra (Southern, 1914)

131106 Aonides oxycephala (Sars, 1862)

131107 Aonides paucibranchiata

Southern, 1914

4 2 4 2

129914 Chaetopterus variopedatus

(Renier, 1804)

129972 Monticellina dorsobranchialis

(Kirkegaard, 1959)

129745 Macrochaeta clavicornis

(M. Sars, 1835)

129892 Mediomastus fragilis Rasmussen, 1973

129220 Notomastus Sars, 1850 1 1

130327 Praxillura longissima Arwidsson, 1906 1

130309 Microclymene tricirrata Arwidsson, 1906

130291 Euclymene droebachiensis

(Sars, 1872)

130980 Scalibregma inflatum Rathke, 1843

129472 Polygordius Schneider, 1868 1 3 10 1 10

130544 Owenia fusiformis Delle Chiaje, 1844

131574 Trichobranchus glacialis

Malmgren, 1866

131474 Amphitrite cirrata (Müller, 1771 in 1776)

131480 Amphitritides gracilis (Grube, 1860)

131516 Pista cristata (Müller, 1776) 1

154972 Pista lornensis (Pearson, 1969)

196385 Polycirrus norvegicus Wollebaek, 1912 1

131009 Hydroides norvegicus Gunnerus, 1768

560033 Spirobranchus lamarcki (Quatrefages, 1866) 3 2 1

555935 Spirobranchus triqueter (Linnaeus, 1758)

137570 Tubificoides amplivasatus

(Erséus, 1975)

2038 Enchytraeidae Vejdovský, 1879 1 1

Crustacea

1078 Ostracoda Latreille, 1802 1

102882 Monoculodes carinatus (Bate, 1857) 1

102918 Pontocrates arenarius (Bate, 1858) 1 5 1

103008 Parapleustes bicuspis (Krøyer, 1838) 7 1

102981 Metaphoxus fultoni (Scott, 1890) 3 1 1

46



#1 PL34

#2 PL34

#3 PL34

#4 PL34

#5

102611 Lysianassa plumosa Boeck, 1871

148560 Socarnes erythrophthalmus

Robertson, 1892

15 8 4 4 12

102132 Atylus vedlomensis (Bate & Westwood, 1862)

1 1

1

102135 Dexamine spinosa (Montagu, 1813)

102137 Guernea (Guernea) coalita

(Norman, 1868)

2

101669 Cheirocratus Norman, 1867 females

101470 Leptocheirus Zaddach, 1884 1 1

102036 Leptocheirus hirsutimanus

(Bate, 1862)

102039 Leptocheirus pectinatus (Norman, 1869) 4 2 6 6

102043 Microdeutopus anomalus

(Rathke, 1843)

148603 Monocorophium sextonae

(Crawford, 1937)

101864 Phtisica marina Slabber, 1769 2

531364 Animoceradocus semiserratus

(Bate, 1862)

6 2 6 2 7

118842 Conilera cylindracea (Montagu, 1804)

1133 Tanaidacea Dana, 1849 1

136443 Leptognathia paramanca

Lang, 1958

1137 Cumacea Krøyer, 1846 1

106791 Processidae Ortmann, 1896 juv.

106687 Paguroidea Latreille, 1802 juvs

107150 Galathea intermedia Liljeborg, 1851

106837 Pisidia Leach, 1820 juv.

107327 Inachus dorsettensis (Pennant, 1777)

107318 Eurynome aspera (Pennant, 1777)

110467 Vaunthompsonia cristata

Bate, 1858

118426 Cymodoce Leach, 1814

Mollusca

234208 Testudinalia testudinalis

(O. F. Müller, 1776)

140199 Leptochiton asellus (Gmelin, 1791)

140201 Leptochiton cancellatus (Sowerby, 1840) 3 3 1

140132 Callochiton septemvalvis

(Montagu, 1803)

139959 Emarginula fissura (Linnaeus, 1758)

605961 Parthenina decussata (Montagu, 1803)

151894 Euspira nitida (Donovan, 1804)

140744 Philine aperta (Linnaeus, 1767) juvs

140640 Onchidoris muricata (O. F. Müller, 1776)

140589 Nucula nitidosa Winckworth, 1930

140590 Nucula nucleus (Linnaeus, 1758)

140472 Musculus discors (Linnaeus, 1767)

47



#1 PL34

#2 PL34

#3 PL34

#4 PL34

#5

140467 Modiolus modiolus (Linnaeus, 1758) juvs 1 2

140235 Limaria hians (Gmelin, 1791)

140283 Lucinoma borealis (Linnaeus, 1767)

141662 Thyasira flexuosa (Montagu, 1803)

140161 Kellia suborbicularis (Montagu, 1803)

345281 Kurtiella bidentata (Montagu, 1803)

146952 Tellimya ferruginosa (Montagu, 1808)

147021 Moerella donacina (Linnaeus, 1758)

181343 Parvicardium pinnulatum

(Conrad, 1831)

1

1 1

139012 Parvicardium scabrum (Philippi, 1844) juvs 1 1 1 1

138159 Spisula Gray, 1837 broken

141577 Arcopagia crassa (Pennant, 1777)

147022 Moerella pygmaea (Lovén, 1846) 2 8

140873 Gari tellinella (Lamarck, 1818) 1 1 3

138636 Dosinia Scopoli, 1777

141911 Dosinia exoleta (Linnaeus, 1758) 6 8 5 1 4

138645 Tapes Megerle von Mühlfeld, 1811

juvs

181364 Venerupis corrugata (Gmelin, 1791)

745846 Polititapes rhomboides (Pennant, 1777)

141909 Clausinella fasciata (da Costa, 1778) 1 1 2 5 1

141929 Timoclea ovata (Pennant, 1777) 1 1

140103 Hiatella arctica (Linnaeus, 1767) 1

141651 Thracia villosiuscula (MacGillivray, 1827) 1 4 5 1

Echinodermata

123080 Asteroidea de Blainville, 1830 juvs

123084 Ophiuroidea Gray, 1840 juvs 18 4 23 8 3

125131 Ophiothrix fragilis (Abildgaard, in O.F. Müller, 1789)

4

1 1 1

125064 Amphipholis squamata (Delle Chiaje, 1828) 4 5 1

123082 Echinoidea Leske, 1778 juvs 1 2

124273 Echinocyamus pusillus (O.F. Müller, 1776)

124418 Spatangus purpureus O.F. Müller, 1776 1

123479 Cucumaria de Blainville, 1830 juv

124685 Neopentadactyla mixta (Östergren, 1898) Deichmann, 1944

1

124465 Leptosynapta inhaerens

(O.F. Müller, 1776) juvs

Tunicata

103718 Ascidiella aspersa (Müller, 1776)

103719 Ascidiella scabra (Müller, 1776)

Phoronida

128545 Phoronis Wright, 1856

Pisces

48



#1 PL34

#2 PL34

#3 PL34

#4 PL34

#5

126928 Pomatoschistus minutus

(Pallas, 1770)

Number of Taxa

31 32 34 26 31

Number of individuals

105 111 142 91 83

% total maerl 92 93 92 93 92

Mean 92.4

% live maerl 7 8 7 8 10

Mean 8


HI22#1

HI22#2

HI22#3

HI22#4

HI22#5

LBN#1

LBN#2

LBN#3

LBN#4

LBN#5

Cnidaria

283798 Cerianthus lloydii 1 1 1 1

100665 Edwardsiidae

Platyhelminthes

793 Platyhelminthes 1

Nemertea

152391 Nemertea 1 1 1 1 2

Nematoda

799 Nematoda 3 2 1 8 2 1 3 2 3 1

Sipuncula

2032 Golfingiidae

410724 Golfingia vulgaris

136060 Nephasoma minutum

Annelida

130707 Pisione remota

939 Polynoidae 3 1 4

130770 Harmothoe impar 1 1 1 1


1


130816 Malmgreniella lunulata 1 1 1 2

130599 Pholoe baltica 2 1 1 1 1 1

130601 Pholoe inornata 2 1

931 Phyllodocidae

130616 Eteone longa


130631 Eulalia mustela

129446 Eumida

130641 Eumida bahusiensis 1 1

130644 Eumida sanguinea 2

129296 Glycera 1

130123 Glycera lapidum 1 2

49


HI22#1

HI22#2

HI22#3

HI22#4

HI22#5

LBN#1

LBN#2

LBN#3

LBN#4

LBN#5

131083 Ephesiella peripatus 1 1 1


1

3

152249 Psamathe fusca 1 2 2 1

130185 Nereimyra punctata 1

131288 Eurysyllis tuberculata 1 1

157583 Syllis cornuta

131458 Syllis variegata

335151 Trypanosyllis coeliaca 2 1 1 6

131379 Sphaerosyllis bulbosa

130417 Platynereis dumerilii 1 1

129838 Pareurythoe borealis 1

130467 Nothria conchylega 1

742232 Lysidice unicornis 1 1 1 2 1

607402 Hilbigneris gracilis 1 2 1



130585 Paradoneis lyra

131106 Aonides oxycephala


1 1 1

1 2


1


1


129892 Mediomastus fragilis 2 1 1 2 3

129220 Notomastus

130327 Praxillura longissima

130309 Microclymene tricirrata


1

130980 Scalibregma inflatum 2 1 1 1

129472 Polygordius 2 6 3 1 1

130544 Owenia fusiformis 1 1


1 1 1 3

131474 Amphitrite cirrata 1

131480 Amphitritides gracilis

131516 Pista cristata 1 1 1 1

154972 Pista lornensis 1 2 1

196385 Polycirrus norvegicus 1 1

131009 Hydroides norvegicus 1 2 1 2 3 1 2

560033 Spirobranchus lamarcki 1 7 3 1

555935 Spirobranchus triqueter 1 1 2 3 2

137570 Tubificoides

50


HI22#1

HI22#2

HI22#3

HI22#4

HI22#5

LBN#1

LBN#2

LBN#3

LBN#4

LBN#5

amplivasatus

2038 Enchytraeidae

Crustacea

1078 Ostracoda

102882 Monoculodes carinatus

102918 Pontocrates arenarius

103008 Parapleustes bicuspis 1

102981 Metaphoxus fultoni 1 1

102611 Lysianassa plumosa 1 3 1 2 1 5


1 2 10

1

102132 Atylus vedlomensis 4 2

102135 Dexamine spinosa


101669 Cheirocratus

101470 Leptocheirus


1

102039 Leptocheirus pectinatus 1 4 2


1

1 2


101864 Phtisica marina


1

2

118842 Conilera cylindracea

1133 Tanaidacea


1

1137 Cumacea

106791 Processidae 1

106687 Paguroidea 1

107150 Galathea intermedia 3 6 6 6

106837 Pisidia 1

107327 Inachus dorsettensis 1

107318 Eurynome aspera 1


3

1

3

118426 Cymodoce

Mollusca


1

1

140199 Leptochiton asellus 1 1 1

140201 Leptochiton cancellatus 3 1 1 2


1

139959 Emarginula fissura 1

51


HI22#1

HI22#2

HI22#3

HI22#4

HI22#5

LBN#1

LBN#2

LBN#3

LBN#4

LBN#5

605961 Parthenina decussata

151894 Euspira nitida

140744 Philine aperta 1 1 2 3

140640 Onchidoris muricata

140589 Nucula nitidosa 1

140590 Nucula nucleus 1

140472 Musculus discors 1

140467 Modiolus modiolus 1 2 1

140235 Limaria hians 1

140283 Lucinoma borealis 1

141662 Thyasira flexuosa 1

140161 Kellia suborbicularis 1

345281 Kurtiella bidentata

146952 Tellimya ferruginosa

147021 Moerella donacina 1


3

2

2

139012 Parvicardium scabrum 1 1 1 2 1

138159 Spisula

141577 Arcopagia crassa 3 1 1

147022 Moerella pygmaea 1

140873 Gari tellinella 1

138636 Dosinia 18 9

141911 Dosinia exoleta 4 1 1 3 4 4 4

138645 Tapes 12 5 7 4

181364 Venerupis corrugata 1 1

745846 Polititapes rhomboides 2

141909 Clausinella fasciata 1 4 3 3 1 1 2 1 1

141929 Timoclea ovata

140103 Hiatella arctica

141651 Thracia villosiuscula 3 5 2 1 2

Echinodermata

123080 Asteroidea

123084 Ophiuroidea 4 4 15 6 8 1 1 1

125131 Ophiothrix fragilis 1 4 5 3 3 13

125064 Amphipholis squamata 22 1 1 4 1 3 6 1 3

123082 Echinoidea 1 1 1 3 1 1 3

124273 Echinocyamus pusillus 1 1 1 2

124418 Spatangus purpureus 1 1 1

123479 Cucumaria

124685 Neopentadactyla mixta


1

Tunicata

52


HI22#1

HI22#2

HI22#3

HI22#4

HI22#5

LBN#1

LBN#2

LBN#3

LBN#4

LBN#5

103718 Ascidiella aspersa

103719 Ascidiella scabra 1

Phoronida

128545 Phoronis 4

Pisces


1

Number of Taxa 21 24 28 24 27 24 26 32 35 34

Number of individuals 35 92 59 50 46 45 57 71 54 77

% total maerl 92 93 93 94 92 95 95 94 94 95

Mean 92.8 94.6

% live maerl 2 1 2 1 1 12 13 12 14 10

Mean 1.4 12.2


HI02#1

HI02#2

HI02#3

HI02#4

HI02#5

HI04#1

HI04#2

HI04#3

HI04#4

HI04#5

Cnidaria

283798 Cerianthus lloydii 2 1

100665 Edwardsiidae 1

Platyhelminthes

793 Platyhelminthes

Nemertea

152391 Nemertea 1 9 1

Nematoda

799 Nematoda 76 36 20 36 3 19 35 25 17 32

Sipuncula

2032 Golfingiidae

410724 Golfingia vulgaris 1

136060 Nephasoma minutum

Annelida

130707 Pisione remota 1 1

939 Polynoidae 2 2 1

130770 Harmothoe impar 1 1 1


1

2

2

2


130816 Malmgreniella lunulata 1 3 1 3 1 2

130599 Pholoe baltica 1 1 2 3 2

130601 Pholoe inornata 1 2 1 2 2 1

931 Phyllodocidae

130616 Eteone longa 1


1

130631 Eulalia mustela 1

53


HI02#1

HI02#2

HI02#3

HI02#4

HI02#5

HI04#1

HI04#2

HI04#3

HI04#4

HI04#5

129446 Eumida 1 1

130641 Eumida bahusiensis

130644 Eumida sanguinea

129296 Glycera 1

130123 Glycera lapidum 1 1 1 1 2 2 1 1

131083 Ephesiella peripatus


1

152249 Psamathe fusca 1

130185 Nereimyra punctata

131288 Eurysyllis tuberculata

157583 Syllis cornuta

131458 Syllis variegata

335151 Trypanosyllis coeliaca 1 2 1 1 1 1

131379 Sphaerosyllis bulbosa 15 3 4 5 1 2 7 2 6

130417 Platynereis dumerilii 1

129838 Pareurythoe borealis

130467 Nothria conchylega

742232 Lysidice unicornis 3 1 1

607402 Hilbigneris gracilis 1 1


5 1

3


130585 Paradoneis lyra 1

131106 Aonides oxycephala 1 1


30 8 4 3 6 8 6 4 5 3




1 2

129892 Mediomastus fragilis 1 2 1 1 5 2

129220 Notomastus 3 1 1

130327 Praxillura longissima

130309 Microclymene tricirrata 1 3


130980 Scalibregma inflatum 1 1 1 2

129472 Polygordius 1 2 1 3 2 1 1 5

130544 Owenia fusiformis


1

131474 Amphitrite cirrata

131480 Amphitritides gracilis 1

131516 Pista cristata 2 1

54


HI02#1

HI02#2

HI02#3

HI02#4

HI02#5

HI04#1

HI04#2

HI04#3

HI04#4

HI04#5

154972 Pista lornensis

196385 Polycirrus norvegicus 2 1 1 1 1 2 1

131009 Hydroides norvegicus 1 1 1 1 1 2

560033 Spirobranchus lamarcki 5 1

555935 Spirobranchus triqueter 1 1 12

137570 Tubificoides amplivasatus

3

1

2038 Enchytraeidae 13 8 3 1 1 2 1

Crustacea

1078 Ostracoda

102882 Monoculodes carinatus

102918 Pontocrates arenarius

103008 Parapleustes bicuspis 1 2 1

102981 Metaphoxus fultoni 1 1 2 1

102611 Lysianassa plumosa 2 1 2 2 1 1


1 9

4

1

1

102132 Atylus vedlomensis 1 2 1

102135 Dexamine spinosa 1


101669 Cheirocratus 1 1

101470 Leptocheirus


102039 Leptocheirus pectinatus



1

101864 Phtisica marina


1

1 2

118842 Conilera cylindracea 1

1133 Tanaidacea


1137 Cumacea

106791 Processidae

106687 Paguroidea 1

107150 Galathea intermedia 2 1

106837 Pisidia

107327 Inachus dorsettensis

107318 Eurynome aspera


118426 Cymodoce 1

Mollusca

55


HI02#1

HI02#2

HI02#3

HI02#4

HI02#5

HI04#1

HI04#2

HI04#3

HI04#4

HI04#5


3 1

1 2 2 1

140199 Leptochiton asellus 3

140201 Leptochiton cancellatus 2 1 4 1 2


139959 Emarginula fissura

605961 Parthenina decussata 1

151894 Euspira nitida 1

140744 Philine aperta

140640 Onchidoris muricata 1

140589 Nucula nitidosa

140590 Nucula nucleus

140472 Musculus discors

140467 Modiolus modiolus

140235 Limaria hians 1

140283 Lucinoma borealis

141662 Thyasira flexuosa

140161 Kellia suborbicularis

345281 Kurtiella bidentata 1

146952 Tellimya ferruginosa 1

147021 Moerella donacina 2


1 1

139012 Parvicardium scabrum 3 2 1 2 2 1

138159 Spisula 1

141577 Arcopagia crassa 2 1

147022 Moerella pygmaea 1 1 1

140873 Gari tellinella 1 1

138636 Dosinia

141911 Dosinia exoleta 1 7 7 2 1 4 4 4 9

138645 Tapes 5 3 8 4 4 4 6 3

181364 Venerupis corrugata

745846 Polititapes rhomboides 1 1 1

141909 Clausinella fasciata 2 2 3 2 4

141929 Timoclea ovata

140103 Hiatella arctica

141651 Thracia villosiuscula 2 1 1 2 2 3

Echinodermata

123080 Asteroidea 1 1

123084 Ophiuroidea 3 13 1 1 3 3 2

125131 Ophiothrix fragilis 2 1 3 4 1 6 1 1

125064 Amphipholis squamata 2 5 14 13 8 5 3 12 1 8

123082 Echinoidea 1 2 1

124273 Echinocyamus pusillus 1 2 3 1

56


HI02#1

HI02#2

HI02#3

HI02#4

HI02#5

HI04#1

HI04#2

HI04#3

HI04#4

HI04#5

124418 Spatangus purpureus

123479 Cucumaria 1

124685 Neopentadactyla mixta


Tunicata

103718 Ascidiella aspersa 1

103719 Ascidiella scabra 1

Phoronida

128545 Phoronis

Pisces


Number of Taxa 32 31 26 22 21 25 38 29 36 31

Number of individuals 183 119 81 90 46 73 101 105 91 98

% total maerl 93 94 92 93 92 98 99 98 98 97

Mean 92.8 98

% live maerl 1 2 0 0 0.5 0 1 0 0 1

Mean 0.7 0.4

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